APHIDS ON

THE WORLD'S TREES

An Identification and Information Guide

R.L. Blackman and V.F. Eastop

Department of Entomology
The Natural History Museum
London
SW7 5BD
UK

CAB INTERNATIONAL
in association with

The Natural History Museum

APHIDS ON THE WORLD'S TREES
An Identification and Information Guide
 CONTENTS

Preface vii

I INTRODUCTION 1
The Association Between Aphids and Trees 1
Life Cycles and Polymorphism 3

II THE TREES AND THEIR APHIDS 14

Notes on the Use of this Section 14
Morphology and Key Characters 15
Host Lists and Keys for Each Tree Genus (in alphabetical order) 21
Key to Polyphagous Tree-dwelling Aphids 532

III THE APHIDS 538

Introduction 538
Systematic Treatment of Aphid Genera (in alphabetical order) 539

IV TECHNIQUES 926
Collecting Methods 926
Preservation and Mounting 927
Labelling and Storage 928

V BIBLIOGRAPHY 930
Regionally Classified Works on Tree-dwelling Aphids 930
List of References 930

VI INDEX TO SPECIES NAMES OF APHIDS 967

VII PHOTOGRAPHS OF SLIDE-MOUNTED APHIDS 987
 PREFACE

One of the first responses that we had to the book Aphids on the World's
Crops, soon after its publication in 1984, was from a colleague in forest
entomology, who pointed out that, despite the title of the work, we had
neglected even to mention a crop that he regarded as rather important, namely
timber. Almost ten years later, we hope that we have now gone some way
towards rectifying that omission.
When we compiled Aphids on the World's Crops (1984) we had good
reason to turn a blind eye to the aphids on timber trees, as we knew that keys
to identify tree-dwelling aphids would be a very different task, much more
complicated than the rather simple keys that we put together ten years ago
to identify the aphids most commonly found on agricultural and horticultural
crops. When we actually got down to the job, we found it necessary to adopt
a far more comprehensive approach than in the earlier work, as we could think
of no justifiable reason for including some aphid species and omitting others.
There was also no way of providing keys to identify aphids that had not been
slide-mounted, as many of the key couplets involve relatively minor differ-
ences between closely related species, requiring microscopic examination. We
think, nevertheless, that non-specialist entomologists will be able to use this
book to identify tree-dwelling aphids at least to genus, and it may then be
possible to confirm the identity of a species by taking into account the
biological information provided in Section III.
Another small problem was to decide what is, and what is not, a tree. We
claim no expertise in this matter, and have relied on standard botanical refer-
ence works, especially Bailey (1949), Uphof (1968), Willis (1973), Boutelje
(1980) and Mabberley (1987). If a genus contains plant species classified as
trees by any or all of these authorities then we have included it. If, as is often
the case, the same genus includes species of shrubby habit, then we have also
generally included such shrubs in the host lists because, in choosing a host plant,
aphids have a lot of regard for its genus but very little regard for its growth habit.

Vll
 Preface

We are grateful to several colleagues who have given permission for the
results of their (as yet) unpublished work to be included; namely B. Agarwala,
S. Akimoto, S. Aoki, S. Barbagallo, P.A. Brown, C.I. Carter, S. Chakrabarti,
R. Danielsson, J. Holman, N. Moran, J.H. Martin, D. Noordam, W.
Quednau, G. Remaudière, J.T. Sorensen, D. Stern, M.B. Stoetzel and D.J.
Voegtlin.
 INTRODUCTION

THE ASSOCIATION BETWEEN APHIDS AND TREES

The known world fauna of aphids (Aphidoidea) consists of 4401 species,
placed in 493 currently accepted genera. Of these, 1758 species in 270 genera
spend all or part of their life feeding on trees. Thus, only 40% of aphid species
but members of 55% of aphid genera occur on trees. The anomaly results from
the numerous members of the evolutionarily recent subfamily Aphidinae
which feed on the evolutionarily more recent herbaceous plants. It will be seen
from Table 1 that all the other major groups of aphids are mostly or even
entirely associated with trees.
The proportion of tree-living aphid species is probably even higher than
indicated, as the unknown hosts of many species are likely to be trees. The
trees most favoured as hosts tend to be the older evolutionary groups such
as Coniferae, Lauraceae, Fagaceae, Betulaceae, Hamamelidaceae, Ulmaceae
and Juglandaceae, and it seems likely that the major groups of aphids differ-
entiated before the appearance of herbaceous plants. Only three groups at the
tribal level live only on herbs; the Saltusaphidini (subfamily Drepanosiphinae)
which live on Cyperaceae and Juncaceae, the Siphini (subfamily Chaito-
phorinae) living on Gramineae, and the Tramini (subfamily Lachninae) living
mostly on roots of Compositae.
Aphids are predominantly a northern temperate group, with remarkably
few species in the tropics. Dixon (1987) postulated that the great diversity of
the tropical forest fauna mitigates against short-lived host-specific insects such
as aphids. Certainly the absence of aphids from many tropical forest trees is
striking, with whole families (e.g. Dipterocarpaceae) seeming almost immune
from attack. The absence of records of aphids from economically important
tropical forest trees such as mahogany (Swietenia mahogoni, Meliaceae) and
rosewood (Dalbergia nigra, Leguminosae) can hardly be due to negligence by
collectors, and suggests that aphids really do not occur on such trees. We

1
2 Introduction

think, however, that the explanation for this can be found in the evolutionary
history of aphids rather than in their present-day host relations or ecology.
Psyllids have similar ecology and host relations to aphids, yet many tropical
trees with few aphids bear a large psyllid fauna. It seems likely to us that
aphids have failed to diversify in the tropics because of one particular,
primitive feature of aphid biology, their cyclical parthenogenesis.
Cyclical parthenogenesis is a very successful way of exploiting the short-
lived growth flushes of temperate plants, but cannot be readily adapted
to tropical conditions. Aphids moving into the tropics simply lose the sexual
phase of the life cycle, and in doing so they lose the potential to evolve
and diversify that is dependent on the recombination of genes. The tropics
may also have acted in this way as a barrier to aphid colonization of southern
temperate regions, which also have very small indigenous aphid faunas.
The occurrence of Neophyllaphis on Podocarpus, Araucaria and related
conifers throughout the southern continents testifies to the age of aphid-tree
relationships, but very little is known about such evolutionarily ancient asso-
ciations. Most ecological and experimental studies of aphid-tree interactions
have concerned introduced species. In Britain, economic damage to spruce by
sporadic outbreaks of Elatobium abietinum has been documented since 1846.
Damage to the more recently introduced Picea sitchensis is particularly severe,
heavy infestations resulting in complete needle loss. Aphid infestations have
been shown to reduce the accretion of wood (e.g. of sycamore; Dixon, 1971a),
and have deleterious effects on tree root growth (e.g. of Tilia; Dixon, 1971b).
However, none of these trees is native to Britain. There are no native British
Picea, sycamore is an introduction from Central Europe, and the common
British lime tree (or linden) is thought to be a hybrid between a native and
an introduced species.
Planted forests of exotic trees cover enormous areas of the globe. There
are more than 5.5 million hectares of planted forests in Brazil, of which at
least 40% are Eucalyptus spp. (Anon., 1985). Pinus radiata occupies only a
small area in its native California but has been widely planted in New Zealand
and elsewhere. During this century many European, oriental and American
species of Pinus were introduced to various parts of Africa and grew aphid-
free for many years. In recent times three aphids, Eulachnus rileyi from
Europe, Cinara cronartii from North America and Pineus boerneri of uncer-
tain origin, have appeared on pines in Africa and caused far greater damage
than they do in Europe or America. Similarly, Cinara cupressi is much more
damaging to Cupressaceae in Africa than in Europe. These exotic conifers may
be growing under stress, and the aphids are certainly without the complex of
natural enemies associated with them in their countries of origin.
Most aphid damage to trees seems to result directly from feeding, either
by removal of sap or wounding of tissue, or in at least some cases by the toxic
effect of saliva. Aphids are rarely recorded as vectors of viruses infecting trees
(Biddle and Tinsley, 1967). Given the astronomical numbers of aphids in the
air and the length of life of trees, there must be strong selection among trees
for resistance to aphid-transmitted viruses. It would be interesting to know
the mechanism of this resistance, and whether it could be transferred to
 Introduction 3

shorter-lived crop plants. Perhaps the energy required to maintain such

defences would be uneconomic for annual or biennial plants.
The problem of how a long-lived plant genotype such as an individual tree
survives, when its herbivores have numerous generations in which to evolve
methods of breaking its defences, is discussed by Whitham (1983), who
showed that there is a similar range of resistance to attack by Pemphigus betas
among different branches of one cottonwood tree, as there is among trees in
a population. He concluded that long-lived plants are mosaics of phenotypic
and/or genetic variability, the genetic differences possibly arising by somatic
mutation (Whitham and Slobodchikoff, 1981).
Alstad and Edmunds (1983) found that the black pine-leaf scale,
Nuculaspis californica, seemed to establish demes with genetic adaptations to
counteract the defence patterns of individual ponderosa pine trees. It is not
known whether any aphids develop such long-term natural associations. Tree-
dwelling aphids, especially those of the large subfamily Drepanosiphinae, tend
to be rather more active insects than the aphids which colonize herbaceous
plants, and may frequently move between trees - although the extent of move-
ment by individual aphids is still largely unknown. Most aphid species in
several other subfamilies alternate annually or biennially between their tree
host and a herbaceous host (see below), and therefore cannot develop
genotype-specific associations, unless of course they were to return to the same
tree year after year.

LIFE CYCLES AND POLYMORPHISM

Aphid life cycles can be quite complicated and involve a succession of morpho-
logically different forms (morphs) of the same species. The complexity - and
the terminology created to describe it - can be daunting to the non-specialist.
Rather than add to the pages of descriptive text already available on aphid life
cycles (e.g. Hille Ris Lambers, 1966e; Blackman, 1974; Dixon, 1985; Miyazaki,
1987), we will merely summarize the essential features, avoiding jargon as much
as possible, and use diagrams (Figs 1-7) to illustrate typical life cycles of tree-
dwelling aphids. Some unavoidable additional terminology - for example that
needed to describe adelgid morphs and life cycles - can be picked up by study-
ing the life cycle diagrams. The essential features of aphid life cycles are:

1. The various families and subfamilies of Aphidoidea each have life cycles
with characteristic features, indicating that they have evolved independently.
2. A complete life cycle (that is, a holocycle) typically consists of one genera-
tion of sexual morphs (sexuales) and several generations in which only parthe-
nogenetic females are produced. This phenomenon of cyclical parthenogenesis
is a basic, primitive feature of aphid biology.
3. In the more primitive families, Adelgidae and Phylloxeridae, both sexual and
parthenogenetic females are oviparous, but in the Aphididae parthenogenetic
females always give birth to live young; in Aphididae the parthenogenetic
4 Introduction

Table 1. Some features of the biology, host associations and life cycles of Aphidoidea. The
bracketed figures are total numbers of genera/total numbers of species : numbers of genera
known to live on trees/numbers of species known to live on trees. The classification follows
that used in B & E (1984) rather than that of Heie (1980), which raises most subfamilies to
family level. Table 3 (p. 539) compares the two classifications.
1 ADELGIDAE (2/49:2/49) alternation from galls on Picea to other Pinaceae, holarctic
1.1 Pineini (1/21:1/21) Picea to Pinus,
1.2 Adelgini (1/28:1/28) Picea to Larix, Pseudotsuga, etc.

2 PHYLLOXERIDAE (8/75: 7/64) Dicotyledons, holarctic

2.1 Phylloxerinini (1/7:1/7) Salicaceae
2.2 Phylloxerini (7/68:7/57) mostly Juglandaceae and Fagaceae, a few spp. on
Ulmaceae, Rosaceae and Vitaceae

APHIDIDAE
3 Pemphiginae (48/319:47/204) dwarf arostrate sexuales, oviparae produce only one egg
3.1 Eriosomatini (13/96 :13/58) galls or pseudogalls on Ulmaceae to various
angiosperms, often on roots, with ants; holarctic
3.2 Pemphigini (18/168 :17/108) holarctic
3.2.1 Pemphigina (6/78 :6/49) galls on Populus to roots of various herbs, more
rarely trees or aerial parts of herbs
3.2.2 Prociphilina (12/90:7/59) pseudogalls on various dicot trees to roots of
Coniferae, more rarely other plants
3.3 Fordini (17/55:17/38) alternation from galls on Anacardiaceae
3.3.1 Fordina (12/41:12/26) Pistacia, mostly to grass roots with ants, mostly
Mediterranean and East Asia, 1 American genus
3.3.2 Melaphidina (5/14:5/12) Rhus, mostly to mosses, mostly oriental, 1 American
genus
4 Hormaphidinae (45/183:42/144) host alternating, with small rostrate sexuales
4.1 Cerataphidini (13/91:10/57) galls on Styrax to Gramineae (esp. Bambuseae),
Palmaceae, Zingiberaceae, where often with ants; oriental
4.2 Hormaphidini (4/11:4/11) galls on Hamamelis to Betula, holarctic
4.3 Nipponaphidini (28/81:28/76) galls on Distylium to Lauraceae, Fagaceae, etc., where
often with ants; oriental
5 Phloeomyzinae (1 ?+) Populus, alate sexuales; all viviparae apterous; holarctic
6 Thelaxinae (3/12:3/12) Betulaceae, Fagaceae, Juglandaceae, often with ants; small
apterous sexuales; holarctic
7 Anoeciinae (2/34:2/15)
7.1 Aiceonini (1/14:1/9) Lauraceae; alate males; Asia
7.2 Anoeciini (1/20:1/6) alternation, Cornus to roots of Gramineae; sexuparae producing
small apterous males
8 Mindarinae (1/5:1/5) Pinaceae; Picea and Abies, few generations per year, holarctic
9 Drepanosiphinae (88/530:74/382) free-living on leaves, mostly on trees; fundatrices often
alate, oviparae apterous, many without ants
9.1 Drepanosiphini (7/45:6/37) Aceraceae; all viviparae alate; holarctic
9.2 Phyllaphidini (72/410:68/345) many on Fagaceae and Betulaceae, some on
Lauraceae, Annonaceae, Magnoliaceae, Ulmaceae, Juglandaceae, Combretaceae,
Burseraceae, Rosaceae, Leguminosae, bamboo; often all viviparae alate
 Introduction 5

Table 1. (cont.)
9.3 Saltusaphidini (9/75:0/0) Cyperaceae and Juncaceae; apterous viviparae common;
holarctic
10 Chaitophorinae (11 /159:6/138) free-living on leaves and shoots, apterous viviparae
common, often with ants
10.1 Chaitophorini (6/138:6/138) holarctic
10.1.1 Chaitophorina (3/91:3/91) Salicaceae
10.1.2 Periphyllina (3/47:3/47) Aceraceae
10.2 Siphini (5/21:0/0) Gramineae; holarctic
11 Greenideinae (18/154:13/87) dicot trees, esp. Fagaceae, mostly East Asia
11.1 Greenideini (9/132:7/77) oviparae often alate
11.2 Cervaphidini (9/22:6/10) mostly southern hemisphere; 4 genera in Australia, 1 in
South America
12 Aphidinae (244/2517:59/299) fundatrices and oviparae large, males apterous or alate;
worldwide but mostly holarctic
12.1 Pterocommatini (5/49:5/37) Salicaceae, without host alternation; holarctic
12.2 Aphidini (25/665:14/90) alternation in many genera but many species are
monoecious; often with ants
12.2.1 Aphidina (20/585:9/67) mostly Rosidae and Asteridae
12.2.2 Rhopalosiphina (5/80:5/23) alternation, Rosaceae to Gramineae and
Cyperaceae
12.3 Macrosiphini (214/1803:40/222) alternation in many genera but many species are
monoecious on herbs, often without ants
13 Lachninae (20/361:14/299) fundatrices and oviparae large, males alate or apterous
(sometimes small); often with ants
13.1 Lachnini (9/57:8/50) mostly Fagaceae and Rosaceae; holarctic, esp. central and
eastern palaearctic
13.2 Cinarini (6/269:5/248) Coniferae; holarctic, most numerous in nearctic
13.3 Tramini (5/35:1/1) roots, mostly Compositae; palaearctic, esp. central and eastern

females are therefore termed viviparae, and the sexual females are distin-
guished as oviparae.
4. The more complex life cycles involve host alternation; the technical term
is heteroecy. In heteroecious aphids, the sexuales mate and fertilized eggs are
laid on a tree or shrub, the primary host, but a regular migration occurs
at some stage in the life cycle to another, totally unrelated plant, which may
be herbaceous or woody - the secondary host. On the secondary host, only
parthenogenetic generations (exules) occur, and a return migration to the
primary host is needed before the next sexual generation.
5. Because host alternation has evolved several times independently in
Aphidoidea, there are important differences at the family and subfamily levels
in the way in which it is achieved (see Figs 1-4). It may occur as part of a one-
year cycle (this happens in all Aphidinae, Hormaphidinae, Pemphigini,
Eriosomatini), or the complete cycle may take two years (Adelgidae, Fig. 1;
Fordina, Fig. 2).
6 Introduction

6. The great majority of aphids go through both the sexual and partheno-
genetic phases of their life cycle on one host plant, or on a small range of
closely-related plants. The technical term for this is monoecy. Drepanosi-
phinae, Chaitophorinae, Greenideinae and Lachninae do not have host
alternation; all species in these subfamilies are monoecious. Some examples
of monoecious life cycles are depicted in Figs 5, 6 and 7. Monoecious aphids
generally have fewer morphs, and there are smaller differences between
morphs, than in heteroecious aphids, although there may be considerable
seasonal variation.
7. Some aphids have lost the sexual part of the life cycle; that is, they are
anholocyclic. Some species are entirely anholocyclic and have no known sexual
morphs (e.g. Tuberolachnus salignus, Pineus boerneri), while others may be
anholocyclic in warmer climates and holocyclic in cold temperate regions
(e.g. Eulachnus rileyi). Populations of certain species maintain the options of
both sexual and parthenogenetic reproduction in mild climates, by producing
sexuales while at the same time continuing to produce parthenogenetic females
(e.g. many Greenideinae). Anholocyclic populations of heteroecious aphids
lose their link with the primary host and live all year reproducing partheno-
genetically on secondary host plants.
 Introduction 7

Fig. 1. Life cycle of Adelges laricis, an example of the 2-year heteroecious cycle of Adelgidae (partly
based on Carter, 1971). Stippled sector is time spent on primary host. Fundatrices hatch in autumn and
the young larvae (1) overwinter on spruce twigs. They feed at the bud scale bases in early spring (2),
inducing a gall (3) in late May-June, which opens in June-July. The alate gallicolae (4) fly to larch and
lay eggs on the needles (5). The larvae hatching from these eggs, the 'neosistens' stage (6),
overwinters on larch twigs, becoming the adult sistens (plural sistentes) in mid-April (7). The
progrediens larvae (8) which hatch from the eggs of the sistens stage develop on new larch shoots,
becoming either sexuparae (9) or apterous progredientes (9a). The sexuparae migrate to spruce in
May-June and lay eggs which develop as sexual males and females (10). The fertilized eggs laid by
the sexual females develop into fundatrix larvae, completing the holocycle. The apterous progredientes
give rise to one or more summer generations on larch, producing copious wax and honeydew.
8 Introduction

Fig. 2. Life cycle of Baizongia pistaciae, an example of the 2-year heteroecious cycle of Fordina in the
Mediterranean area and southwest Asia (based mainly on Wertheim, 1954). Stippled sector is time
spent on primary host. Fundatrix larvae (1) hatch from eggs on Pistacia in late March-April and feed on
young shoots causing bending (2). They later move to a leaflet which becomes folded and eventually
develops into a large pod-like gall (3). Two generations of apterae occur in the gall before alatae start
to be produced. The gall splits open near its apex in late October-November, releasing large numbers of
emigrant alatae (4), which found colonies of apterous viviparae on roots of grasses (5). Alate sexuparae
(6) are produced by these colonies in March-April and return to Pistacia, where they give birth to very
small sexual females (oviparae) and males (7) on the bark. The oviparae each mature a single egg
which is not laid, the ovipara dying with the egg still inside her. The egg (9) does not hatch until the
following spring.
 9

Fig. 3. Life cycle of Thecabius affinis, an example of the life cycle of a heteroecious member of the
Pemphiginae (based on Mordvilko, 1935; Roberti, 1938; Danielsson, 1976). Stippled sector is time
spent on the primary host. In late spring, the fundatrix (1) inhabits a small gall formed by folding one
edge of a poplar (Populus nigra) leaf. Her progeny leave this gall and feed along the mid-rib of another
leaf, causing it to fold in half longitudinally (2), become roughened and blistered on the outer surface
and eventually turn red. The progeny of the fundatrix developing inside this gall all become emigrant
alatae (3), which emerge in June-July and give rise to waxy colonies of apterous exules (4) at stem
bases and on runners of Ranunculus repens. In September, alate sexuparae (5) are produced which fly
to trunks of poplar and give birth to the very small, arostrate males and oviparae (6). (Colonies may
also persist on the secondary host and alate viviparae may be produced that found new colonies on
Ranunculus.) The sexuales develop rapidly to adult and mate on the poplar bark (7). The oviparae each
lay a single egg (8) which does not hatch until the following spring.
10 Introduction

Fig. 4. Life cycle of Aphis fabae, as an example of a heteroecious member of the Aphidinae (after
Blackman, 1974). Stippled sector is time spent on the primary host. The fundatrix (1) develops from an
overwintering egg and founds a colony on new growth of Euonymus europaeus in spring, which gives
rise to emigrant alatae (2) in May-June. Successive generations occur on various herbaceous secondary
hosts through spring and summer (3); alate viviparae are produced as the colonies become larger and
more crowded, and these fly to found fresh colonies on other plants. In autumn, in response to
decreasing daylength, the colonies on the secondary hosts produce alate gynoparae (4) and alate males
(5), which migrate back to Euonymus. The progeny of the alate gynoparae are apterous sexual females,
the oviparae (6), which mature on the spindle leaves just before they fall, and mate with males arriving
independently from secondary hosts. The oviparae lay overwintering eggs (7) in the bud axils.
 Introduction 11

Fig. 5. Life cycle of Periphyllus testudinaceus, holocyclic and monoecious on Acer spp., e.g. A.
campestre (based mainly on Essig and Abernathy, 1952). The first instar fundatrix (1) hatches early in
February and develops on swelling buds and young growth of Acer, becoming adult (2) in March. She
has numerous progeny, both apterous and alate. Some of the alatae (3) fly to other maple trees and
produce further generations of apterous (4) and alate (5) viviparae. Other progeny of the fundatrix, both
apterous (6) and alate (7), remain on the same tree; they produce specialized aestivating first instar
nymphs ('dimorphs') with foliate hairs, which spend the summer scattered on both upper and lower
sides of leaves (8). By mid-June, the only individuals to be found are the dimorphs. Eventually, in
September, these start to develop into adult sexuparae, which have a distinct cruciform dark dorsal
patch (9). In late October-November their progeny, small alate males and apterous sexual females
(oviparae), reach maturity and mate (10). Overwintering eggs (11) are laid on branches in bark crevices
and on twigs near winter buds.
12 Introduction

Fig. 6. Life cycle of Drepanosiphum platanoidis on Acer pseudoplatanus, as an example of the

monoecious, holocyclic annual life cycle of Drepanosiphinae (based largely on work by Dixon, 1971-83).
Eggs hatch in late March and the first instar fundatrix (1) feeds on swelling buds and young spring
growth, maturing in April. The fundatrix is alate (2). Her progeny develop into paler alate adults with
few or no dorsal abdominal markings, which enter into a reproductive diapause lasting until September
(not all Drepanosiphinae aestivate in this way). The pale green summer alatae (3) adopt a spaced-out
distribution when settled on the undersides of leaves, but move around and fly readily when disturbed.
Parthenogenetic reproduction resumes in September, with a generation of alate sexuparae (4), usually
with well-developed dorsal abdominal cross-bands. These give rise to the sexuales (5); rather narrow-
bodied males with conspicuous dark dorsal markings, and large brown sexual females (oviparae) - the
only apterous morph in this species. Overwintering eggs (6) are laid on the twigs.
 Introduction 13

Fig. 7. Life cycle of Cinara schwarzii on Pinus ponderosa in western USA, as an example of the
monoecious, holocyclic annual life cycle of Cinarini (after Palmer, 1926). First instar fundatrices hatch
in late March (1) and feed on the twigs, developing to adult (2) in April. Large, ant-attended colonies of
viviparae, including both apterae (3) and alatae (4), develop on the bark of twigs and small branches.
In some species of Cinara alatae occur only in June-July, but in C. schwarzii they are found from May
through to September. Alate males (5) and apterous oviparae (6) occur in September-October. (Males of
some species of Cinara are apterous, and oviparae often have a conspicuous perianal ring of white
wax.) The shiny black eggs (wax-covered in some species) are laid in rows along the needles (7).
 THE TREES AND THEIR APHIDS

NOTES ON THE USE OF THIS SECTION

The aphids occurring on any one agricultural crop are mostly in different
genera so that they can be identified using a relatively simple key (Blackman
and Eastop, 1984). Identification of tree-dwelling aphids present far greater
problems, because aphids and trees have mostly coevolved, with the result that
closely-related aphid species frequently occur on closely-related tree species.
Specificity is rarely absolute, however, so that keying the aphids on each
species of tree is impractical, and most keys are at the level of the tree genus.
A key to the aphids on Abies, for example, thus has to distinguish between
many aphid species within the genus Cinara.
The user of these keys will therefore need to make microscope slide whole
mounts of the aphids to be identified. We recommend that Canada balsam
mounts are prepared as they are of proven permanence, whereas the water-
based Berlese mountant once widely used for aphid preparations is liable to
degenerate in time. A simple procedure for preparing balsam mounts, that of
Martin (1983), is given on p. 928. It does not involve staining; phase contrast
microscopy is recommended. Many of the keys involve measurements of parts
of the aphid such as antennal segments with a micrometer eyepiece or other
measuring device. Correct and accurate calibration of the measuring device
is obviously very important.
This section contains:

1. Essential information about the morphology of tree-living aphids, the

characters and abbreviations used in the keys and some warnings about the
pitfalls that await anyone starting to identify aphids.
2. A list of the plant families with tree genera from which aphids are recorded
(Table 2).

14
 The Trees and Their Aphids 15

3. The tree genera arranged in alphabetical order with host lists of the aphids
recorded from each tree species, followed by or including keys to the aphids.
The arrangement varies slightly according to the number of aphid species
and their degree of specificity. If numerous aphid species in one genus are
involved - Cinara on pines or Eriosoma on elms for example - then we have
taken advantage of any specificity shown by the aphids to limit the number
of species that have to be discriminated in any one key. The very large aphid
faunas of some tree genera - Quercus and Salix for example - are divided up
in a preliminary 'master' key, to avoid long and cumbersome keys with 100
or more couplets. Species which are keyed on the basis of published descrip-
tions and have not been seen by the authors are indicated by an asterisk (*).
4. This section ends with a key to the 23 species of polyphagous aphids that
may each be found on various tree genera. Many tree genera, particularly
those in the tropics and southern hemisphere, have no aphids specific to them,
but are sometimes colonized by such polyphagous aphids, particularly when
there is vigorous new growth. Most of these aphid species are also found on
herbaceous plants, and some of them are important pests of agricultural crops.
A few of the species included are not strictly polyphagous, but tend to limit
their feeding to both woody and herbaceous members of one plant family; for
example Aphis craccivora on many genera of Leguminosae.

MORPHOLOGY AND KEY CHARACTERS

The morphological terminology is illustrated for a generalized aphid in Fig. 8;
for more detailed information see Miyazaki (1987). The parameters measured
are as in Ilharco and van Harten (1987), except that measurement of body
length (BL) does not include the CAUDA. Four commonly used measure-
ments are illustrated in Fig. 9. The following abbreviations are used:

BL Body length (excluding any projecting CAUDA)

ANT I, ANT II, etc. Antennal segments I, II, etc or their lengths
ANT PT Terminal process of last antennal segment, or its
length (Fig. 9A)
PT/BASE RATIO Length of ANT PT divided by length of basal
part of last segment including primary rhinarium
(Fig. 9A)
R IV+V Last rostral segment, or its length (in fact the
complex formed by R IV and R V, which are
sometimes referred to separately; Fig. 9B)
HT I and HT II First and second segments of hind tarsus, or
their lengths (Fig. 9C)
ABD TERG 1, 2, etc. Abdominal tergites 1, 2, etc.
 16 The Trees and Their Aphids

Fig. 8. Morphology of a generalized adult female alate viviparous aphid, with parts named in accordance
with terminology used in keys. In this example the media of the forewing is twice-branched.

SIPH Siphunculus, or its length

CAUDA CAUDA, or its length (Fig. 9D)

Most keys will only work for the particular morph or morphs for which
they were intended. Wherever possible the morph keyed is the adult apterous
vivipara (aptera for short), but there are plenty of exceptions. Many
 The Trees and Their Aphids 17

Fig. 9. Measurements frequently required in the keys: A, length of last antennal segment (base and
processus terminalis); B, length of last segment of rostrum (R IV+V); C, length of second segment of
hind tarsus (HT II); D, length of CAUDA.

Drepanosiphinae do not produce any apterous viviparae, and in the case of

gall-forming Pemphiginae and Hormaphidinae the most convenient morph to
collect in large numbers and identify is the alata that emerges from the
gall. In Adelgidae on secondary hosts, the overwintering first instar sistens
('neosistens') has useful recognition features, whereas the identification of
summer populations is very difficult.
Keys will not work if applied to immature aphids, and in several groups,
especially when the adult CAUDA is broad and rounded as in Lachninae,
Greenideinae and many Pemphiginae, it can be difficult to distinguish between
late instar immatures and apterous adults. If the form of the adult CAUDA
is not distinctive, then the presence of rudimentary gonopophyses, and of
fully-formed and clearly defined anal and genital plates, can provide recogni-
tion features for adults, but even these are not wholly reliable. To be confident
about the recognition of adults in difficult cases it may be necessary to
examine a number of specimens of different sizes from the same colony, in
order to establish what are the particular features of the adult insect in that
species. In any case collection and examination of a large sample should be
the general rule, because identifications should always be based on an
examination of the fullest possible range of variation.
One point that we wish to stress is that these keys are only one part of
18 The Trees and Their Aphids

the identification process. They may give the user a possible name for the
specimens, but no reliance should be placed on this alone. They are based on
existing host plant records, and it is quite possible that aphids will be collected
that are not in the key because they have not previously been recorded from
the plant in question. This is particularly likely in places where the aphid fauna
is less well known. More confidence in the identification may be gained if the
information given in Section III under that name also fits the specimens, but
even then one should proceed with caution; perhaps take the further step of
comparing your aphids with museum specimens identified by a recognized
authority, or consult a more detailed published description. When correct
identification is essential - in biological control work for example - it is
always advisable to consult a taxonomie specialist, who has learnt by experi-
ence many of the pitfalls involved!

Table 2. Tree genera with aphids on them, arranged by families [The nomenclature follows
Brummitt (1992), with some additional subfamily divisions from Willis (1973) and Mabberley
(1987).]___________________________________________

Aceraceae: Acer
Anacardiaceae
Anacardieae: Anacardium, Buchanania, Mangifera
Pistaciaceae (Rhoideae): Harpephyllum, Lannea, Loxostylis, Ozoroa, Pistacia, Rhus, Schinus
Spondieae: Sclerocarya, Spondias
Annonaceae: Annona, Xylopia
Apocynaceae: Apocynum, Carissa, Conopharyngia, Rauwolfia, Tabernaemontana
Aquifoliaceae: Ilex
Araliaceae: Aralia, Cussonia, Eleutherococcus (= Acanthopanax), Kalopanax, Panax, Polyscias,
Pseudopanax, Schefflera
Araucariaceae: Agathis, Araucaria
Asclepiadaceae: Calotropis, Margaretta

Betulaceae: Alnus, Betula

Bignoniaceae: Catalpa, Jacaranda, Kigelia, Markhomia, Radermachera, Spathodea, Tabebuia,
Tecoma
Bixaceae: Bixa, Scolopia
Bombacaceae: Bombax, Chorisia, Ochroma
Boraginaceae: Cordia, Ehretia, Tournefortia
Buddlejaceae: Buddleja
Burseraceae: Canarium, Commiphora

Caprifoliaceae: Sambucus
Caryocaraceae: Caryocar
Casuarinaceae: Casuarina
Cecropiaceae: see Moraceae
Celastraceae: Celastrus, Euonymus, Maytenus
Cercidiphyllaceae: Cercidiphyllum
Chrysobalanaceae: Chrysobalanus, Parinaria
Combretaceae: Anogeissus, Combretum, Terminalia
Compositae: Piptocarpha, Vernonia
Cornaceae: Cornus
 The Trees and Their Aphids 19

Table 2. (cont.)
Corylaceae: Carpinus, Corylus, Ostrya
Cupressaceae: Callitris, Calocedrus, Chamaecyparis, X Cupressocyparis, Cupressus, Juniperus,
Pilgerodendron, Tetraclinis, Thuja, Widdringtonia
Cyatheaceae: Cyathea (= Alsophila)
Cycadaceae: Cycas
Cyrillaceae: Cyrilla

Dipterocarpaceae: Dipterocarpus, Monotes

Ebenaceae: Diospyros, Euclea

Elaeagnaceae: Elaeagnus, Hippophae
Elaeocarpaceae: Aristotelia, Elaeocarpus
Ericaceae: Arbutus
Escalloniaceae: Escallonia
Eucommiaceae: Eucommia
Euphorbiaceae
Acalyphoideae: Alchornea, Macaranga, Mallotus, Ricinus
Crotonoideae: Aleurites, Croton
Euphorbioideae: Euphorbia, Sapium
Phyllanthoideae: Bischofia, Bridelia, Glochidion, Phyllanthus, Securinega
Eupteleaceae: Euptelea

Fagaceae: Castanea, Castanopsis (= Shiia), Fagus, Lithocarpus, Nothofagus, Quercus

Flacourtiaceae: Casearia, Doryalis, Scolopia

Gramineae: Arundinaria, Bambusa, Dendrocalamus, Gigantochloa, Phyllostachys, Sasa

Greyiaceae: Greyia
Guttiferae: Calophyllum, Cratoxylum, Garcinia, Harungana, Mammea, Mesua

Hamamelidaceae: Corylopsis, Distylium, Hamamelis, Liquidambar

Hernandiaceae: Hernandia
Hippocastanaceae: Aesculus
Hymenocardiae: Hymenocardia

Juglandaceae: Carya, Engelhardtia, Juglans, Platycarya

Labiatae: Hyptis
Lauraceae: Actinodaphne, Cinnamomum, Laurus, Lindera, Litsea, Machilus, Nectandra,
Neolitsea, Ocotea, Persea, Phoebe, Sassafras, Syncarpia, Umbellularia
Lecythidaceae: Barringtonia
Leguminosae
Caesalpinioideae: Brachystegia, Caesalpinia, Cassia, Ceratonia, Cercis, Cynometra,
Dimorphandra, Gleditsia, Julbemardia, Peltophorum, Schotia, Tamarindus
Mimosoideae: Acacia, Albizia, Inga, Leucaena, Mimosa, Pithecellobium, Prosopis
Papilionideae: Aganope (= Ostryoderris), Andira, Castanospermum, Dalbergia, Laburnum,
Lonchocarpus, Maackia, Millettia, Mundulea, Pterocarpus, Robinia, Sesbania,
Sophora, Virgilia
Loganiaceae: Strychnos
Lythraceae: Duabanga, Lagerstroemia
20 The Trees and Their Aphids

Table 2. (cont.)
Magnoliaceae: Liriodendron, Magnolia, Michelia
Malpighiaceae: Banisteriopsis, Malpighia
Malvaceae: Hibiscus, Plagianthus, Thespesia
Meliaceae: Cedrela, Dysoxylum, Khaya, Melia, Toona
Melianthaceae: Bersama
Memecylaceae: Memecylon
Moraceae: Artocarpus, Brosimum, Broussonetia, Cecropia, Cudronia, Ficus, Morus, Streblus
Moringaceae: Moringa
Myricaceae: Myrica
Mysinaceae: Rapanea
Myrtaceae
Leptospermoideae: Eucalyptus, Leptospermum, Melaleuca, Metrosideros, Syncarpia, Tristania
Myrtoideae: Eugenia, Myrtus, Psidium, Syzigium

Naucleaceae: Nauclea, Sarcocephalus

Nyctaginaceae: Pisonia, Torrubia
Nyssaceae: Nyssa

Oleaceae: Fraxinus, Ligustrum, Olea, Syringa

Onagraceae: Fuchsia

Palmae: Ancistrophyllum, Arenga, Areca, Calamus, Cocos, Daemonorops, Livistona, Rhapis,

Salacca
Pandanaceae: Pandanus
Pinaceae: Abies, Cedrus, Keteleeria, Larix, Picea, Pinus, Pseudotsuga, Tsuga
Pittosporaceae: Hymenosporum, Pittosporum
Platanaceae: Platanus
Podocarpaceae: Podocarpus
Polgonaceae: Calligonum
Proteaceae: Grevillea, Macadamia

Rhamnaceae: Berchemia, Hovenia, Rhamnus, Ziziphus

Rosaceae
Rosoideae: Hagenia
Prunoideae: Amygdalus, Prunus
Pyroideae (Pomoidea): Amelanchier, Crataegus, Eriobotrya, Heteromeles, Malus, Photinia,
Pyrus, Sorbus
Rubiaceae
Cinchonoideae: Cinchona, Craterispermum, Crossopteryx, Gardenia, Hymenodictyon, Kraussia,
Morinda, Psychotria, Randia, Rothmannia, Tocoyena
Guettardoideae: Guettarda
Rutaceae: Calodendrum, Citrus, Euodia, Murraya, Phellodendron, Tectea, Toddalia, Vepris,
Zanthozylum

Sabiaceae: Meliosoma
Salicaceae: Chosenia, Populus, Salix
Sapindaceae: Dodonaea, Koelreuteria, Pappea, Sapindus
Sapotaceae: Malacantha, Mimusops, Pouteria
Scrophulariaceae: Paulownia
 The Trees and Their Aphids 21

Table 2. (cont.)
Simaroubaceae: Aiianthus
Sonneratiaceae: Duabanga (see Lythraceae)
Staphyleaceae: Euscaphis, Staphylea
Sterculiaceae: Brachychiton, Dombeya, Guazuma, (Theobroma)
Stilaginaceae: Antidesma
Styracaceae: Sinojackia, Styrax
Symplocaceae: Symplocos

Tamaricaceae: Tamarix
Taxaceae: Taxus
Taxodiaceae: Cryptomeria, Metasequoia, Sequoia, Sequoiadendron, Taxodium
Theaceae: Camellia, Schima, Stewartia
Tiliaceae: Corchorus, Grewia, Microcos, Schoutenia, Tilia
Trochodendraceae: Euptelia, Trochodendron

Ulmaceae: Aphananthe, Celtis, Hemiptelea, Planara, Trema, Ulmus, Zelkova

Umbeiliferae: Steganotaenia
Urticaceae: Laportea

Verbenaceae: Citharexylum, Holmskioldia, Premna, Tectona, Vitex

Vochysiaceae: Qualea

Xanthophyllaceae: Xanthophyllum

Zamiaceae: Encephalartos
Zygophyllaceae: Guaiacum

HOST LISTS AND KEYS FOR EACH TREE GENUS

(in alphabetical order)

ABIES True Firs Pinaceae

About 25 Cinara species are described from Abies, but many of the North
American species are unrecorded since their original descriptions or little
known. Abies is the secondary host for all species of Adelges subgenus
Dreyfusia, which are undoubtedly the most economically important group,
migrating from cone-like galls on their primary host Picea, or reproducing
parthenogenetically on Abies throughout the year.
22 Abies

Host Plant List

Abies alba Adeiges merken, nordmannianae,
(Silver Fir) pectinatae, piceae
Cinara confinis, costata, indica,
pectinatae
Mindarus abietinus
Prociphilus americanus, bumeliae,
fraxini, oriens
A. amabilis Adelges pectinatae, piceae
(Cascade Fir, Pacific Silver Fir) Cinara chinookiana, occidentalis
A. balsamea Adelges nordmannianae, pectinatae,
(Balsam Fir) piceae
Cinara confinis, curvipes
Mindarus abietinus
Prociphilus americanus, bumeliae
A. bornmuelleriana Adelges nordmannianae
Cinara confinis, pectinatae
Elatobium abietinum, sp. nr
momii
Mindarus abietinus
A. cephalonica Adelges piceae, prelli
(Greek Fir) Cinara confinis
Mindarus abietinus
A. cilicia Adelges piceae
Cinara confinis, curvipes
Mindarus abietinus
A. concolor Adelges pectinatae, piceae
(White Fir) Cinara alacra, curtihirsuta,
curvipes, gentneri, grande,
kiusa, occidentalis, osborni,
radicivora, schuhi, sonata,
zoarcbursara
Essigella pergandei
Mindarus abietinus
A. delavayi Prociphilus fraxini
A. faberi Adelges glandulae
A. faxoniana Adelges piceae
Mindarus abietinus
A. firma Adelges piceae
(Momi Fir) Cinara longipennis, matsumurana,
todocola
 Abies 23

Elatobium momii
Mindarus abietinus, japonicus
A. forrestii Adelges piceae
A. fraseri Adelges piceae
(Southern Balsam Fir) Mindarus abietinus
A. grandis Adelges nordmannianae, pectinatae,
(Grand Fir) piceae
Cinara confinis, curvipes, gentneri,
occidentalis, pacifica, sonata
Mindarus abietinus, victoria
A. holophylla Cinara longipennis, smaragdina
(Needle Fir) Mindarus japonicus
A. homolepis Cinara todocola
(Nikko Fir) Mindarus abietinus
A. koreana Adeiges pectinatae, piceae
A. kosteri Elatobium abietinum
A. lasiocarpa Adelges pectinatae, piceae
(Subalpine Fir) Cinara chinookiana, confinis,
curvipes, minuta, occidentalis,
osborni
Mindarus abietinus
A. magnifica (incl. var. shastensis) Cinara curvipes, setulosa, sonata
A. mariesii Cinara hattorii
A. mayriana (see A. sachalinensis var. mayriana)
A. nebrodensis Adelges nebrodensis, piceae
Mindarus abietinus
A. nephrolepis Adelges pectinatae
[Prociphilus bumeliaeformis
Mordvilko ex Shaposhnikov,
1955]
Cinara smaragdina
A. nobilis Adelges nordmannianae, piceae
Cinara confinis, curvipes
A. nordmanniana Adelges merken, nordmannianae,
pectinatae, piceae, prelli
Cinara confinis
Elatobium abietinum, sp. nr
momii
Mindarus abietinus
Prociphilus fraxini
24 Abies

A. numidica Cinara pectinatae

Mindarus abietinus
A. pindrow Adelges joshii, knucheli, piceae
pindrowi
Cinara confmis, Cinara sp.
Mindarus japonicus
A. pinsapo Cinara curvipes
A. procera Adelges pectinatae
Prociphilus americanus
A. religiosa Cinara curvipes
A. sachalinensis Cinara confinis, hattorii,
(Saghalin Fir) longipennis, matsumurana,
todocola
Mindarus abietinus, japonicus
A. sachalinensis var. mayriana Adelges pectinatae ssp. ishiharai,
todomatsui
Cinara hattorii, longipennis,
matsumurana, todocola
Mindarus japonicus
Prociphilus oriens
A. sibirica Adelges pectinatae
Cinara confinis
Mindarus abietinus
A. spectabilis Adelges knucheli
(Himalayan Fir) Cinara sp.
A. veitchii Adelges pectinatae, todomatsui
Cinara pectinatae, todocola
Mindarus abietinus
Abies spp. Cinara abietihabitans, [tistaensis]
Pineus abietinus

Key to Abies-feeding aphids

All the Abies-feeding aphids are keyed together, as there is little evidence that
any of them are specific to particular Abies species. Except where otherwise
stated, the key should be applied only to adult apterous females. With Cinara
species, care must be taken that the individuals examined are not immature
(see p. 17), as this can lead to erroneous identifications. Cinara species not
seen by us (identified with an *) are keyed according to Hottes (1960c),
and identifications of these species in particular should be treated very
circumspectly.
Colonies of Prociphilus spp. in white wax wool on roots of Abies cannot
 Abies 25

Fig. 10. Dorsal (right) and ventral (left) views of adult sistens of A, Pineus abietinus and B, Adeiges
nordmannianae.

be satisfactorily keyed to species, and identification of Adelges to species

requires the neosistens stage to be available.

1 Adult apterae with very short antennae, less than 0.1 of body length, with
at most 3 segments. Small insects (BL less than 1.2mm) producing white
wax wool. Always oviparous, adults having a distinct chitinous ovipositor
(Adelgidae) .................................................. 2
- Adult apterae with 5- or 6-segmented antennae, at least 0.2 of body length.
BL 1.3-7.8 mm. With or without wax wool. Viviparous in spring and
summer (Aphididae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Body of adult aptera broadly pear-shaped to almost spherical. Cephalo-
prothoracic shield entire, uniformly pigmented. Abdomen with 4 pairs of
evident spiracles. Wax pores discontiguous (Fig. 10A) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus abietinus
- Body of adult aptera pear-shaped or oval. Head and prothorax not
uniformly pigmented. Abdomen with 5 pairs of evident spiracles. Wax
pores contiguous (Fig. 10B) . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges spp.
(see key to resting-stage first instar larvae of Adelges spp. at end of main
key)
3 SIPH tubular. ANT PT/BASE equal to or greater than 1.0 . . . . . . . 4
- SIPH broadly conical, pore-like or absent. ANT PT/BASE less than 1.0
............................................................. 5
4 ANT PT/BASE 1.6 or less . . . . . . . . . . . . . . . . . . . Elatobium abietinum
- ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . Elatobium momii
26 Abies

Fig. 11. SIPH of aptera of A, Cinara curvipes and B, Cinara confinis.

5 SIPH present as pores on hairy cones which are usually pigmented

(Fig.11) ..................................................... 6
- SIPH either absent or present as pores without pigmented conical bases
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
6 (Couplets 6-27 refer to apterous viviparae)
Maximum diameter of base of SIPH cone less than 0.3 mm, or less than
3 times the diameter of the SIPH aperture . . . . . . . . . . . . . . . . . . . . . . . 7
- Maximum diameter of base of SIPH cone (or the pigmented area
incorporating it) more than 0.3 mm, or more than 3 times the diameter
of the SIPH aperture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
7 HT I at least half as long as HT II (Fig. 12A). SIPH cones pale ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pectinatae
- HT I less than half as long as HT II; if nearly half as long, then SIPH
cones dark ................................................... 8
8 HT II at least 4 times longer than the very short, almost triangular
HT I (Fig. 12B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara occidentalis
- HT II 2.1-3.5 times longer than HT I . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 SIPH cones very small and pale. Longest hairs on ANT III more than
twice as long as width of segment . . . . . . . . . . . . Cinara matsumurana
- SIPH cones dark. Longest hairs on ANT III less than twice width of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 BL less than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara minuta*
- BL more than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . Cinara zoarcbursara*
 Abies 27

Fig. 12. HT I and II of A, Cinara pectinatae and B, Cinara occidentalis.

11 SIPH cones with hairs of two sizes, the longest hairs at least twice as long,
and clearly thicker than, the shortest hairs (Fig. 11B) . . . . . . . . . . . . 12
- SIPH cones with hairs varying in length by less than a factor of 2
(Fig.11A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
12 SIPH cones pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara setulosa
- SIPH cones dark ............................................ 13
13 Pigmented bases of SIPH cones very extensive, with no clear limits,
extending anteriorly and posteriorly to join pigmented lateral areas on
other abdominal segments (Fig. 13A) . . . . . . . . . . . . . . . . Cinara osborni
- Pigmented bases of SIPH cones less extensive, delimited, restricted to
ABD TERG 6, or 5 + 6, any pigmented lateral areas on other segments
being separate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Length of sclerotized part of stylet groove less than 1.8 mm. ANT V about
equal in length to, or shorter than, ANT VI (including PT) . . . . . . 15
- Length of sclerotized part of stylet groove (Fig. 74, p. 292) greater than
1.8mm. ANT V clearly longer than ANT VI including PT . . . . . . . 17
15 BL greater than 4 mm. Hind tibiae conspicuously bicoloured, with yellow
region on basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara alacra*
28 Abies

Fig. 13. SIPH and surrounding sclerites of aptera of A, Cinara osborni and B, Cinara sonata.

- BL less than 4mm. Hind tibiae uniformly pigmented, usually dark...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Length of sclerotized part of stylet groove less than 1.3mm. Hairs on
ABD TERG 1-7 without pigmented bases . . . . . . . Cinara chinookiana
- Length of sclerotized part of stylet groove greater than 1.3 mm. Hairs on
ABD TERG 1-7 mostly arising from small pigmented sclerites of various
sizes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara todocola
17 Length of sclerotized part of stylet groove 2.9-3.8 mm ..............
............................................... Cinara radicivora
- Length of sclerotized part of stylet groove less than 2.9 mm . . . . . . 18
18 Antennae conspicuously bicoloured, each segment with a broad black
distal band contrasting with a pale base, ANT IV and V being about half
black and half pale. Pigmented bases of SIPH cones of irregular shape,
usually with a deep notch on the inner margin (Fig. 13B) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara sonata
- Antennae not conspicuously bicoloured, although segments often gradually
darker towards apices. Pigmented bases of SIPH cones more-or-less
rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 Dorsal length of HT I greater than its width at base (Fig. 14A) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara tenuipes
- Dorsal length of HT I less than its width at base (Fig. 14B) . . . . . . 20
20 ANT III at least 1.1 x ANT IV and V together, and more than 3 x ANT
VI (incl. PT) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara longipennis
 Abies 29

Fig. 14. HT I of A, Cinara tenuipes and B, Cinara confinis.

- ANT III about equal to or a little shorter than ANT IV and V together,
and less than 3 x ANT VI (incl. PT) . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 R IV over 3 times longer than R V . . . . . . . . . . Cinara abietihabitans*
- R IV less than 3 times longer than R V . . . . . . . . . . . . . . . . . . . . . . . . 22
22 BL 3.8-7.8 mm. ANT PT/BASE less than 0.4 . . . . . . . Cinara confinis
- BL 3.2-4.4 mm. ANT PT/BASE equal to or greater than 0.4. (Alatae with
forewing membrane between Cu1a and Culb fuscous) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara hattorii
23 All tibiae black or very dark brown, although sometimes with a slightly
less dark dusky brown section on basal half . . . . . . . . . . . . . . . . . . . . 24
- Fore and mid-tibiae pale yellow to amber, dark at apices if at all, and hind
tibiae if mainly dark then at least with paler section at base . . . . . . 26
24 BL greater than 4.8mm. Hairs on hind tibiae clearly longer than width
of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara grande*
- BL less than 4.8 mm. Hairs on hind tibiae longer or shorter than width
of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 BL less than 3.2mm. ANT III with up to 5 rhinaria. Hairs on hind
tibiae mainly longer than width of tibia at its midpoint . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara schuhi*
- BL greater than 3.2 mm. ANT III with or without rhinaria. Hairs on hind
tibiae shorter than width of tibia at its midpoint . . . . . . Cinara kiusa*
26 Coxae dark brown to black . . . . . . . . . . . . . . . . . . . . . . . Cinara curvipes
- Coxae yellow to amber . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
30 Abies

27 BL less than 4 mm. Hairs on SIPH cones sparse, totalling about 20, with
basal part of cone nearly devoid of hairs . . . . . . . . Cinara curtihirsuta
- BL more than 5 mm. Hairs on SIPH cones very numerous all over
pigmented area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara gentneri
28 Abdomen without wax pore plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
- Abdomen with segmental wax pore plates, at least marginally .... 30
29 Body oval. Hairs on ANT III longer than basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pacifica*
- Body elongate, about 3 times longer than its greatest width in dorsal view.
Hairs on ANT III shorter than basal diameter of segment . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Essigella pergandei*
30 Antennae of aptera shorter than rostrum. Hairs on antennae and legs
long. (On roots; cannot be keyed to species) . . . . . . . Prociphilus spp.
- Antennae of aptera longer than rostrum. Hairs on antennae and legs
short. (On needles) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31 ANT III of alata with 6-12 squarish to almost circular rhinaria (Fig. 15A)
............................................... Mindarus victoria
- ANT III of alata with 12-27 narrow, tranversely elongate rhinaria ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 ANT III of alata with rhinaria in a single row, mostly extending across
complete width of segment (Fig. 15B) . . . . . . . . . . . Mindarus abietinus
- ANT III of alata with rhinaria not aligned in a single row, many of those
on basal half not extending across full width of segment (Fig. 15C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mindarus japonicus

Key to resting-stage (neosistens) first instar larvae of Adelges spp. on Abies

Separation of the spring and summer apterous adult stages of Adelges spp.
(mainly subgenus Dreyfusia) on Abies is very difficult because of their
complex polymorphism. The key given here is based on the work of several
authors (notably Eichhorn, 1958), and relies on the lesser degree of variability
of the more sclerotized first instar larvae of the sistens morph (the neosistens
stage) which, in all species except A. pectinatae and A. pindrowi, is normally
found on the bark of the stem and branches rather than on the needles, and
is the most frequent stage at which aestivation or hibernation occurs. Except
where otherwise stated, all characters used are those of the neosistens. Slide-
mounted preparations are essential for this key.

1 Spinal and pleural sclerites fused on all segments, and spinopleural

sclerites also fused to marginal sclerites of ABD TERG 6-8 (Fig. 16A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges pectinatae/pindrowi
 Abies 31

Fig. 15. ANT III of alata of A, Mindarus victoria, B, M. abietinus and C, M. japonicus.

- Spinal and pleural sclerites of mesothorax, metathorax and ABD TERG

1-5 not fused (sometimes touching) (Fig. 16B) . . . . . . . . . . . . . . . . . . . 2
2 Pleural sclerites without wax glands. Spinal sclerites with well-developed
wax pore plates along inner margins (Fig. 16B) . . . . . . . . . . . . . . . . . . . 3
- Pleural sclerites with wax glands. Spinal sclerites with wax pore plates
variably developed, located centrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3 Wax pore plates on inner margins of meso- and metathoracic spinal
sclerites containing numerous small, mostly rounded, pits (Fig. 16C),
arranged in 2-4 areas of rather rounded shape, the most central area
containing 7-12 pits. Total number of pits in the central areas only of
32 Abies

Fig. 16. Adelges on Abies (neosistens stage except where otherwise indicated). A, dorsal sclerites of
A. pectinatae; B, dorsal sclerites of Adelges (Dreyfusia) sp., showing also the five central areas referred
to in couplets 3 and 4; C, spinal sclerites on meso- and metathorax of A. nordmannianae; D, same for
A. piceae (after Pschorn-Walcher and Zwölfer, 1958); E, second instar sistens of A. prelli (after
Francke-Grosmann, 1937a); F, wax pore plate on spinal sclerite of mesonotum of A. knucheli (after
Schneider-Orelli and Schneider, 1954); 6, thoracic sclerites and wax pores of A. joshii; H, the same for
A. todomatsui (after Inouye, 1953).

the spinal wax pore plates of the meso- and metathorax plus ABD TERG
1-3 (i.e. a total of 10 central areas) is 57-104 .....................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges nordmannianae
- Wax pore plates on inner margins of meso- and metathoracic spinal
sclerites divided into 3-4 angular or rounded areas (Fig. 16D), the most
 Abies 33

central area containing 2-8 relatively large pits of irregular shape, rounded
or polygonal. Total number of pits in central areas of spinal wax pore
plates of meso- and metathorax plus ABD TERG 1-3 (i.e. 10 central areas)
is 18-63 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Central areas of wax pore plates on meso- and metathoracic spinal
sclerites usually more-or-less rounded, each with 3-8 pits. Total number
of pits on the 10 central areas (as defined in couplet 3) is in range 26-63
(usually more than 40) . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges merkeri
- Central areas of wax pore plates on meso- and metathoracic spinal
sclerites often more-or-less triangular, each with 3-6 pits. Total number
of pits on the 10 central areas is usually less than 40 (range 18-59)
............................................................. 5
5 Neosistens strongly sclerotized. 2nd instar sistens with a series of dark
conical humps on the head, thorax and 1st to 3rd (or 4th) abdominal
segments, diminishing in size posteriorly (Fig. 16E) . . . . . . Adelges prelli
- Neosistens variably sclerotized, but usually with rather weak sclerotization
(at least in European populations). 2nd instar sistens without pronounced
sclerotic humps on head, thorax and anterior abdominal segments . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges piceae
6 Dorsal wax pore plates of mesothorax, metathorax and abdomen com-
prising groups of 15-25 small rounded facets . . . . . . . Adelges glandulae
- Dorsal wax pore plates either much larger and taking up most of the area
of the sclerite, or much smaller and composed of small groups of no more
than 7 facets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Spinal, pleural and marginal sclerites strongly sclerotized, those of the
mesothorax, metathorax and abdomen each almost fully occupied by
a large wax pore plate composed of 2-5 segments separated by grooves,
each segment having a reticulate structure (Fig. 16F) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges knucheli
- Spinal, pleural and marginal sclerites weakly sclerotized, with wax pore
plates on mesothorax, metathorax and abdomen much smaller, comprising
only 2-7 grouped or loosely-connected facets, and without reticulate
structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Marginal sclerites of meso- and metathorax each with a single wax pore
plate comprising a compact group of 4-7 facets (Fig. 16G) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges joshii
- Marginal sclerites of meso- and metathorax each containing 2 distinct wax
pore plates, each usually with 4 facets (Fig. 16H) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges todomatsui
34 Acacia

ACACIA Leguminosae

Acacia spp. Aphis craccivora, fabae, gossypii,

spiraecola
Aulacorthum solani
Myzus cymbalariae, persicae
(Use key to polyphagous aphids, p. 532)

ACANTHOPANAX = ELEUTHEROCOCCUS

ACER Maples and Sycamores Aceraceae

There are several aphid genera which are virtually specific to Acer, and within
these genera there is a high degree of monophagy, although a few species are
able to colonize several Acer species.
The first key takes adult aphid specimens colonizing any Acer to species,
with the exception of the two largest genera, Periphyllus (35 spp.) and
Drepanaphis (20 spp.). The species of Acer are then listed in alphabetical
order, with a list of the aphids recorded from them followed by keys, where
necessary, to the species of Periphyllus and/or Drepanaphis.
Finally, a key is given to Periphyllus spp. described from unidentified
species of Acer, together with the seven most polyphagous (oligophagous)
species of this genus (acericola, bulgaricus, californiensis, kuwanaii, lyro-
pictus, testudinaceus and villosi).
To identify an Acer-feeding aphid, use the main key first, then if you know
the species of Acer refer to the alphabetical list to see whether your answer is
appropriate, using the subsidiary keys to Periphyllus and Drepanaphis if
necessary. If you have a species of Periphyllus from an unidentified Acer, or
one that does not appear in the alphabetical list, use the final key. The final
key may also help if your Periphyllus does not seem to be one of those
previously recorded from your species of Acer.
Except where otherwise stated, all the keys can be applied to both apterous
and alate viviparous females, but are unlikely to work for fundatrices,
oviparae and males.
 Acer 35

Fig. 17. SIPH of aptera of Stomaphis aceris.

Main key to aphids on Acer

1 PT short and stump-like, if finger-like then PT/BASE no more than
0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE clearly more than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2 SIPH pores placed at apices of broad, pigmented, hairy cones (e.g.
Fig. 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- SIPH absent, or present as small pores not placed on pigmented, hairy
cones ........................................................ 6
3 Rostrum short, less than one quarter of body length . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- Rostrum longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 HT II of aptera less than 1.5 times longer than segment II of fore or mid-
tarsus. ANT VI (including PT) a little shorter than ANT V. Ventral
abdomen without a median longitudinal row of dark patches . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis graffii
- HT II of aptera 2 or more times longer than segment II of fore
or mid-tarsus. ANT VI (including PT) longer than ANT V. Ventral
36 Acer

Fig. 18. A, ANT III of spring migrant alata of Neoprociphilus aceris; antenna of spring migrant alata of
B, Mimeuria ulmiphila and C, Paraprociphilus tessellatus.

abdomen with a median longitudinal row of 5 elongate dark patches

............................................................. 5
5 Dorsal abdomen of aptera with paired large, dark patches on all segments.
HT II usually a little shorter than greatest width of dark patch around
SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis aceris
- Dorsal abdomen of aptera without paired large, dark patches. HT II
considerably longer than greatest width of dark patch around SIPH
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis takahashii
6 Antennae of apterae 5- or 6-segmented, about one third of body length.
Spring migrant alatae with rather small, oval secondary rhinaria confined
to basal 0.7 of ANT III (Fig. 18A) . . . . . . . . . . . Neoprociphilus aceris
- Antennae of apterae 5-segmented, less than 0.25 of body length. Spring
migrant alatae with transversely elongate secondary rhinaria extending at
least over ANT III to V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ANT III of aptera (= fundatrix) 1.2-1.9 x ANT V (including PT). ANT
III of spring migrant alata 1.8-2.6 x ANT VI (incl. PT); ANT VI without
secondary rhinaria (except those associated with primary rhinarium)
(Fig. 18B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mimeuria ulmiphila
- ANT III of aptera (= fundatrix) 1.0-1.2 x ANT V. ANT III of spring
migrant alata 1.3-1.7 x ANT VI; ANT VI with or without secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
 Acer 37

Fig. 19. A, CAUDA of Megalophyllaphis sp.; B, aestivating nymph of Periphyllus americanus; C, SIPH of
aptera of Drepanosiphoniella aceris.

8 ANT VI of spring migrant alata with 4-8 secondary rhinaria (Fig. 18C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraprociphilus tessellatus
- ANT VI of spring migrant alata with 0-1 secondary rhinaria . . . . . . . .
....................................... Paraprociphilus mexicanus
9 SIPH generally stump-shaped, those of aptera with an apical zone of
polygonal reticulation, and those of alata generally with more extensive
reticulation. Antennal hairs usually long and conspicuous, mostly much
longer than diameter of segment from which they arise (except in one
North American species on A. glabrum and one East Asian species on
A. rufinerve). Alatae never with dark markings on wings . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Periphyllus spp. (see separate keys)
- SIPH of variable shape, usually without polygonal reticulation, or if with
some reticulation near apices, then adults are all alatae with dark markings
on wings. Antennal hairs generally small and inconspicuous, usually
shorter than segment from which they arise; if there are long hairs then
they are confined to ANT III (with occasionally one also at the base of
ANT IV) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 CAUDA with an elongate knob, the knob more than twice as long as wide
(Fig. 19A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Megalophyllaphis sp(p).
- If CAUDA knobbed then the knob is about as long as wide ..... 11
11 Hairs on front of head and sides of abdomen modified as flattened leaf-
like lamellae (Fig. 19B). (Very small insects, BL less than 1 mm) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . aestivating nymphs of Periphyllus spp.
- Hairs not thus modified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Front of head and at least the sides of the abdomen with long hairs, those
on frons exceeding length of ANT I . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
38 Acer

Fig. 20. Marginal hairs of mesothorax and metathorax of aptera of A, Trichaitophorus aceris (after
Takahashi, 1937a), B, T. japonicus (after Sorin, 1979b) and C, T. koyaensis.

- Hairs all shorter, those of frons much shorter than ANT I . . . . . . . 19

13 SIPH longer than basal width, stump-shaped, conspicuously flared at
apices (Fig. 19C) . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphoniella aceris
- SIPH pale, small, shorter than basal width and not or hardly flared at
apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Dorsal hairs on abdomen as long as marginal hairs . . . . . . . . . . . . . . . .
....................................... Yamatochaitophorus albus
- Dorsal abdominal hairs minute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Antennae of apterae 5- or 6-segmented, as long as or longer than forelegs,
with ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichaitophorus recurvispinus
- Antennae of apterae 4- or 5-segmented, shorter than forelegs, with ANT
PT/BASE less than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 SIPH black. Marginal hairs lanceolate; dilated in middle, with pointed
apices (Fig. 20A) . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichaitophorus aceris
- SIPH pale. Marginal hairs tapering or cylindrical, pointed, rounded or
blunt at their apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 Body elongate oval, about twice as long as wide in dorsal view.
Mesonotum with only one long (posterior) marginal hair on each side
(Fig. 20B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichaitophorus japonicus
- Body broadly oval, about 1.7 times as wide as long in dorsal view.
Mesonotum with 2 pairs of long marginal hairs (Fig. 20C) . . . . . . . 18
18 Antenna about one half of BL. Longest hair on hind tibia about twice
width of tibia at midlength . . . . . . . . . . . . . . Trichaitophorus koyaensis
- Antenna about one third of BL. Longest hair on hind tibia about
equal to width of tibia at midlength . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trichaitophorus aenigmatosus*
 Acer 39

Fig. 21. Forewing of A, Drepanosiphum aceris and B, D. braggii.

19 Adult viviparous females all alate (sometimes brachypterous). CAUDA

shorter than its width at base or, if longer, then clearly constricted and
with a knobbed apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
- Adult viviparae apterous or alate. CAUDA tongue-shaped, longer than
its basal width and never with a knobbed apex . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . see key to polyphagous aphids (p. 532)
20 SIPH tubular, more than 4 times longer than their basal width,
cylindrical, tapering or slightly swollen in middle, without reticulated
apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- SIPH usually flask-shaped, conical or tapering, usually less than 4 times
their basal width or, if longer, then with apical reticulation . . . . . . 28
21 Forewing with a dusky patch at tip between ends of Rs and M (Fig. 21 A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum aceris
- Forewing sometimes with dusky spots at ends of veins, but not with
pigment extending between vein endings . . . . . . . . . . . . . . . . . . . . . . . . 22
22 SIPH tapering gradually from base to apex, rarely with any discernible
swelling. Pterostigma of forewing with a characteristically shaped dark
band, thickened near its proximal end (Fig. 21B). ANT III with less than
12 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum braggii
- SIPH with at least a slight swelling of middle region. Pterostigma of
forewing pale, or with either a uniformly thin dark band or a dark distal
patch. ANT III usually with more than 12 rhinaria . . . . . . . . . . . . . . 23
40 Acer

23 Dorsal abdomen with two broad, dark, transverse bands on segments 4

and 5, that on segment 5 extending laterally to almost touch the marginal
sclerites in front of the SIPH (Fig. 22A). Pterostigma with a small dark
patch at its distal extremity. Often brachypterous . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum dixoni
- Dorsal abdomen either without transverse bars or with more, shorter, bars
not extending to lateral sclerites. Pterostigma without any distinct small
dark patch. Always fully winged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Fore femora more than 1.5 times thicker than mid- and hind femora, and
with a very distinct broad black longitudinal ventral stripe (Fig. 22C).
ANT III with a short black contrasting section near base, as dark as the
apex of the segment . . . . . . . . . . . . . . . . . . . Drepanosiphum oregonense
- Fore femora less than 1.5 times thicker than mid- and hind femora,
sometimes with a dark ventral stripe but this never very conspicuous. ANT
III pale or dusky at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 PT less than 0.7 times as long as ANT III, PT/BASE 6.0-7.0. R IV+V
at least 0.14 mm. BL 3.1-4.3 mm. With or without a black patch in front
of each SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
- PT more than 0.7 times as long as ANT III, PT/BASE 7.5-12.0.
R IV+V less than 0.14mm. With a black patch in front of each SIPH
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
26 Either with no dorsal or lateral abdominal markings (spring/summer) or
with a series of brown-black transverse dorsal bars and lateral patches
including a large black ante-siphuncular patch (autumn) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum platanoidis
- Always with a single black patch just in front of each SIPH, even in
specimens without dorsal abdominal bars (Fig. 22B) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum iranicum
27 SIPH with conspicuously dark tips, even in very pale specimens, and often
wholly dusky or dark. ANT III with 9-15 rhinaria . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum acerinum
- SIPH usually pale, without conspicuously dark tips. ANT III with 15-26
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanosiphum caucasicum
28 SIPH without any reticulation at apices. Forewings either clear or with
pigment only along wing veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
- SIPH with subapical reticulation. Forewings with areas of dark pigment
extending between wing veins, or at least between Rs and media .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
29 SIPH truncate, not swollen on basal half and with a very small flange
(Fig. 22D). Abdomen with only small dorsal processes. Primary rhinarium
on ANT VI without a medium-sized secondary rhinarium proximal to it
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shenawheum minutum
 Acer 41

Fig. 22. A, dorsal abdomen of Drepanosiphum dixoni; B, dorsal abdomen of D. iranicum; C, fore femur
of Drepanosiphum oregonensis; D, SIPH of Shenawheum minutum; E, SIPH of Drepanaphis acerifoliae; F,
forewing of Yamatocallis takagii; 6, forewing of Y. hirayamae; H, forewing of Y. tokyoensis; I, SIPH of
Yamatocallis takagii, J, SIPH of Y. hirayamae.
42 Acer

- SIPH flask-shaped, with swollen base, narrow 'neck' and well-developed

apical flange (Fig. 22E). Abdomen usually with conspicuous pigmented
dorsal processes. Primary rhinarium on ANT VI with a medium-sized
secondary rhinarium proximal to it, and usually another distal to it
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanaphis spp. (see separate keys)
30 SIPH more than 0.6 mm long (Fig. 22I) . . . . . . . . . . . . . . . . . . . . . . . . 31
- SIPH less than 0.5 mm long (Fig. 22J) . . . . . . . . . . . . . . . . . . . . . . . . . 34
31 ANT PT/BASE less than 1.5 . . . . . . . . . . . . . . . . . Yamatocallis sauteri
- ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 ANT III with 12-18 rhinaria. Forewing with Rs and distal branches of M
heavily bordered with black (Fig. 22F) . . . . . . . . . . Yamatocallis takagii
- ANT III with 14-31 rhinaria. Forewing with pigmentation more evenly
distributed, not concentrated along Rs . . . . . . . . . . . . . . . . . . . . . . . . . 33
33 ANT III with 29-31 secondary rhinaria. Primary rhinarium on ANT V
very large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Yamatocallis acericola
- ANT III with 14-28 secondary rhinaria. Primary rhinarium on ANT V
of normal size, about equal in diameter to base of ANT V . . . . . . . . .
......................................... Yamatocallis brevicauda
34 Wing membrane pale, except at bases of veins . . . . . . . . . . . . . . . . . . . . .
........................................... Yamatocallis obscura
- Wings at least partly pigmented between veins . . . . . . . . . . . . . . . . . . 35
35 Forewing mostly pigmented, except for hind margin (Fig. 22H). ANT III
2.4-3.1 times longer than base of ANT VI . . . . . . . . . . . . . . . . . . . . . . . .
......................................... Yamatocallis tokyoensis
- Forewing pigmented on anterior half only (Fig. 22G). ANT III 3.2-5.6
times longer than base of ANT VI . . . . . . . . . Yamatocallis hirayamae

Host Plant List

Acer acuminatum = A. caudatum
A. atlantica Drepanosiphoniella aceris
A. barbatum Drepanaphis carolinensis
Periphyllus americanus
A. buergerianum Megophyllaphis sp.
(Trident Maple) Periphyllus acerihabitans
Yamatocallis hirayamae
A. caesium Periphyllus aceriphaga, caesium,
(Indian Maple) villosi
Trichaitophorus aceris
 Acer 43

Key to Periphyllus species on Acer caesium

1 CAUDA constricted at base. SIPH more than 0.25mm long, almost
cylindrical, pale in aptera and pale or dusky in alata . . . . . . . . . villosi
CAUDA not constricted at base. SIPH less than 0.25 mm long, conical,
dark at least distally in aptera and wholly dark in alata . . . . . . . . . . . 2
2 SIPH 1.0-1.3 times longer than HT II . . . . . . . . . . . . . . . . . . aceriphaga
SIPH more than 1.5 times longer than HT II . . . . . . . . . . . . . . caesium

A. campestre [Aphis aceriella Theobald] = A.

(Hedge Maple) fabae?
[Chaitophorus flavissimus
Mamontova, 1979]
Drepanosiphoniella aceris ssp.
fugans
Drepanosiphum acerinum, dixoni,
platanoidis
Mimeuria ulmiphila
Periphyllus aceris, californiensis,
hirticornis, obscurus,
testudinaceus, vandenboschi,
venetianus, villosi
Stomaphis graffii

Key to Periphyllus species on Acer campestre

1 CAUDA broadly rounded (crescent-shaped), less than half as long as
basal width (Fig. 23A). ANT PT/BASE of aptera 2.0-3.6 (of alata, 1.9-4.1)
............................................................. 2
CAUDA broadly tongue-shaped or with a constriction near base, more
than half as long as its basal width (Figs 23B-D). ANT PT/BASE of
aptera 3.2-6.9 (of alata, 3.7-6.7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Hind tibiae uniformly pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aceris
Hind tibiae at least distally pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Hind tibiae uniformly dark . . . . . . . . . . . . . . . . . . . . . . . . . . californiensis
Hind tibiae with pale middle region, contrasting with dark base and distal
section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . testudinaceus
4 The 2 hairs on base of ANT VI very unequal in length, the longer one
more than 4 times as long as the shorter one, which is less than half length
of base of ANT VI (including primary rhinarial complex) (Fig. 24A)
............................................................. 5
The 2 hairs on base of ANT VI both long and fine, the longer one 1.3-3.0
times the shorter, which is more than half length of base of ANT VI
(Fig.24B) .................................................... 6
44 Acer

Fig. 23. CAUDA of A, Periphyllus testudinaceus, B, P. hirticornis, C, P. obscurus, D, P. lyropictus.

Fig. 24. ANT BASE VI of A, Periphyllus venetianus, B, P. obscurus and C, P. testudinaceus.

5 Length of the shorter of the 2 hairs on base of ANT VI less than basal
diameter of segment. Apterae with long hairs on antennae, legs and dor-
sum all blunt or slightly expanded at their apices, and with SIPH at least
1.5 times as long as their basal widths (Fig. 25A) . . . . . . . . . hirticornis
Length of the shorter of the 2 hairs on base of ANT VI greater than basal
diameter of segment. Apterae with long hairs on antennae, legs and
 Acer 45

Fig. 25. SIPH of A, Periphyllus hirticornis and B, P. venetianus.

dorsum all fine-pointed, and with SIPH about as long as or a little longer
than their basal widths (Fig. 25B) . . . . . . . . . . . . . . . . . . . . . . . venetianus
6 SIPH pale or only slightly dusky, 1.5-2.0 times longer than their basal
widths ................................................... villosi
SIPH dark and only about as long as their basal widths ... obscurus

A. cappadocicum Drepanosiphum iranicum

Periphyllus testudinaceus
A. carpinifolium Yamatocallis acericola
A. caudatum Periphyllus aceriphaga,
garhwalensis, villosi
Trichaitophorus aceris,
? recurvispinus
Yamatochaitophorus albus

Key to Periphyllus species on Acer caudatum

1 CAUDA not constricted at base . . . . . . . . . . . . . . . . . . . . . . . aceriphaga
- CAUDA constricted at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH of alata dark and conical, about as long as its basal width (aptera
not known) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . garhwalensis
SIPH of alata (and aptera) pale, almost cylindrical, more than 1.5 times
longer than basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . villosi

A. circinatum Drepanosiphum oregonense

(Vine Maple) Periphyllus californiensis,
lyropictus, testudinaceus
(Use key to Periphyllus spp. on p. 56)
A. floridanum = A. barbatum
46 Acer

A. formosanum var. coreanum Periphyllus californiensis

A. ginnale Drepanosiphum platanoidis
(Amur Maple) Periphyllus aceris, loricatus,
[takahashii], testudinaceus

Key to Periphyllus species on Acer ginnale

1 Legs wholly pale ......................................... aceris
Legs wholly or partly pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT PT/BASE less than 1.5. Aptera with broad dark bands across all
abdominal tergites, merging intersegmentally . . . . . . . . . . . . . . loricatus
ANT PT/BASE more than 2.5. Aptera with a pattern of discrete sclerites
on dorsum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . testudinaceus

A. glabrum Periphyllus americanus,

(Rocky Mountain Maple) brevispinosus

Key to species on Acer glabrum

1 Longest hairs on antennae more than 2.5 times longer than basal diameter
of ANT III. ANT PT/BASE more than 2.0 . . . . . . . . . . . . americanus
Longest hairs on antennae only a little longer than basal diameter of ANT
III. ANT PT/BASE less than 2.0 . . . . . . . . . . . . . . . . . . . . brevispinosus

A. granatense Drepanosiphum oregonensis

Periphyllus bulgaricus
A. grandidentatum Drepanaphis granovskyi,
(Western Big-toothed Maple) idahoensis, knowltoni, utahensis
Drepanosiphum platanoidis
Periphyllus americanus,
testudinaceus

Key to Drepanaphis species on Acer grandidentatum

1 Fore femora pigmented along their entire lengths, especially dorsally.
Dorsal tubercles on ABD TERG 3 well developed, pigmented, united for
about half their lengths; those on ABD TERG 1 undeveloped and those
on ABD TERG 2 small (Fig. 26A) . . . . . . . . . . . . . . . . . . . . . . knowltoni
Fore femora pale, or dusky at their distal ends only. Dorsal abdominal
tubercles variously developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT PT/BASE more than 8.0. Dorsal abdominal tubercles all pigmented,
those on ABD TERG 3 well developed, dark, united for about half their
lengths; those on ABD TERG 2 conical, about one third as long; those
on ABD TERG 1 small (Fig. 26B) . . . . . . . . . . . . . . . . . . . . . . idahoensis
 Acer 47

- ANT PT/BASE less than 7.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3 Dorsal tubercles on ABD TERG 3 bluntly conical, united at bases, usually
pigmented; those on ABD TERG 1 and 2 undeveloped or small. ANT
PT/BASE 6.0-7.0 (Fig. 26C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . utahensis
- Tubercles on ABD TERG 1, 2 and 3 all small, only 0.5-2.0 x their basal
widths, not united at bases, always pale (Fig.26D). ANT PT/BASE
5.0-6.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . granovskyi

For Periphyllus species use couplet 2 at key on p. 52.

A. griseum Drepanosiphum platanoidis

(Paperbark Maple) Periphyllus testudinaceus
A. hermoneum Drepanosiphoniella aceris
Drepanosiphum oregonense
Periphyllus bulgaricus
A. hyrcanum Periphyllus bulgaricus
A. ibericum Drepanosiphoniella aceris ssp.
caucasica
Periphyllus aceris
A. japonicum Periphyllus californiensis
(Fullmoon Maple) Yamatocallis hirayamae,
tokyoensis
A. laevigatum Periphyllus sp.
(Chinese Maple, Dieng-than)
A. laxiflorum Drepanosiphum platanoidis
Periphyllus testudinaceus
A. macrophyllum Drepanosiphum oregonense
(Big-leaf Maple, Broad-leaf Periphyllus californiensis,
Maple) lyropictus, testudinaceus
(For separation of Periphyllus species see key, p. 56)
A. mandshuricum Periphyllus mandshuricus
A. mayrii Periphyllus kuwanaii
A. miyabei Periphyllus californiensis,
(Miyabei Maple) hokkaidensis
(see key to Periphyllus on A.
mono)
A. mono Periphyllus brevisetosus,
(Japanese or Painted Maple, californiensis, [diacerivorus],
Itaya-kaede) hokkaidensis, kuwanaii, viridis,
[viridis ssp. osugiensis]
Trichaitophorus aceris
Yamatocallis hirayamae, takagii
 48 Acer

A. knowltoni B. idahoensis

C. utahensis D. granovskyi

E. carolinensis F. acerifoliae

G. saccharini H. keshenae

I. sabrinae J. simpsoni

K. parva L. kanzensis
 Acer 49

Key to Periphyllus species on Acer mono

1 ANT PT/BASE more than 3.0. Dorsum of aptera pale . . . . . . . viridis
ANT PT/BASE less than 3.0. Dorsum of aptera with dark transverse bars
or paired sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Hind tibiae of aptera uniformly black. Head with 4-5 pairs of dorsal hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . californiensis
Hind tibiae of aptera at least with central section paler. Head with 6-10
pairs of dorsal hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . kuwanaii
- ANT PT/BASE less than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Longest dorsal cephalic hair 4-4.5 x basal diameter of ANT III . . . . .
.................................................. hokkaidensis*
Longest dorsal cephalis hair about 2.7 x basal diameter of ANT III
................................................... brevisetosus*

A. monspessulanum Drepanosiphoniella aceris, aceris

(Montpellier Maple) ssp. fugans
Drepanosiphum acerinum,
oregonense, platanoidis
Periphyllus bulgaricus, hirticornis,
rhenanus, testudinaceus

Key to Periphyllus species on Acer monspessulanum

1 Hind tibiae with middle part much paler than base or distal section, and
hind femora with pale basal half and contrastingly dark distal half.
Longer of the 2 hairs on ANT BASE VI 21-47 µm long (Fig. 24C). SIPH
of both aptera and alata dark . . . . . . . . . . . . . . . . . . . . . . . testudinaceus
Legs without contrasting pigmentation. Longer of 2 hairs on ANT BASE
VI 45-130 µm long. SIPH of aptera pale . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT PT/BASE more than 4.0. Aptera with most of the long dorsal hairs
pale, blunt or slightly furcate at their apices . . . . . . . . . . . . . hirticornis
- ANT PT/BASE less than 3.5. Long dorsal hairs all fine-pointed, and at
least those on ABD TERG 5-8 are dark-pigmented . . . . . . . . . . . . . . . 3
3 Aptera with long spinal hairs on ABD TERG 5-8 (or 4-8) much darker
than spinal hairs on more anterior tergites. Longer of 2 hairs on ANT
BASE VI 0.8-1.3 x length of ANT BASE VI, and shorter hair
0.25-0.47 x BASE VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rhenanus

Fig. 26. Drepanaphis: anteriolateral view of spinal tubercles on ABD TERG 1-4 of 12 species.
50 Acer

Aptera with long spinal hairs similarly pigmented on all ABD TERG.
Longer of 2 hairs on ANT BASE VI 0.3-0.8 x length of BASE VI, and
shorter hair 0.12-0.27 x BASE VI . . . . . . . . . . . . . . . . . . . . . . bulgaricus

A. negundo Aphis spiraecola see key to

(Box-Elder) Aulacorthum solani polyphagous
Myzus persicae aphids, p. 532
Drepanosiphum braggii,
platanoidis
Paraprociphilus mexicanus
Periphyllus californiensis,
[kuwanaii],
negundinis, [nevskyi,]
testudinaceus
[Stomaphis longirostris?]

Key to Periphyllus species on Acer negundo

1 Tibiae with middle part much paler than base or distal section . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . testudinaceus
Tibiae rather uniformly pigmented, dusky or dark . . . . . . . . . . . . . . . . 2
2 Aptera and alata both with a clear pattern of dorsal dark markings, the
alata with broad transverse bars and the aptera with either bars or paired
spots. Longest hair on base of ANT VI usually more than half as long
as base of ANT VI. Alata with 10-23 rhinaria on ANT III . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . californiensis
Aptera and alata with only faint dorsal abdominal markings. Longest hair
on base of ANT VI always less than half as long as base of ANT VI. Alata
with 3-10 rhinaria on ANT III . . . . . . . . . . . . . . . . . . . . . . . . negundinis

A. nigrum Drepanaphis carolinensis,

(Black Maple, Hard Maple) kanzensis, knowltoni (all
vagrants?)

Key to Drepanaphis species on Acer nigrum

1 Fore femora pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . kanzensis
Fore femora pigmented along entire length, especially dorsally ..... 2
2 ANT PT/BASE more than 8.0. Tubercles on ABD TERG 3 finger-like,
united for more than half their lengths; those on ABD TERG 2 and 4
very small, those on ABD TERG 1 undeveloped (Fig. 26A) . . . . . . . . .
...................................................... knowltoni
- ANT PT/BASE less than 7.0. Tubercles on ABD TERG 3 conical, united
only at base; those on ABD TERG 1, 2 and 4 half as long, subequal,
finger-like (Fig. 26E) ................................. carolinensis
 Acer 51

A. nuttallii Myzus persicae

Periphyllus sp. [negundinis?]
A. oblongum Drepanosiphum oregonense
Periphyllus californiensis,
testudinaceus
(See key to Periphyllus spp. on p. 56)
A. obtusatum Drepanosiphum platanoidis
Periphyllus rhenanus
A. opalus Drepanosiphum oregonensis,
(Italian Maple) platanoidis
Periphyllus testudinaceus,
rhenanus
(See key to Periphyllus spp. on A. monspessulanum)
A. opulifolium Periphyllus bulgaricus
A. orientate Drepanosiphum platanoidis
Periphyllus rhenanus
A. palmatum Periphyllus californiensis,
(Japanese Maple) testudinaceus
Yamatocallis tokyoensis
(For Periphyllus see key p. 56)
A. pennsylvanicus Drepanosiphum platanoidis
(Moosewood)
A. pictum = A. mono
A. platanoides Drepanaphis acerifoliae
(Norway Maple) Drepanosiphum platanoidis
Periphyllus aceris, coracinus,
[kuwanaii], lyropictus,
testudinaceus, [viridis]

Key to Periphyllus species on Acer platanoides

1 CAUDA tongue-shaped, about as long as broad, and slightly constricted
basally (knobbed) (Fig. 23D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . lyropictus
CAUDA broadly rounded or crescent-shaped, clearly shorter than its
basal width (Fig. 23A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Hind tibia with pale middle region contrasting with dark base and distal
section. Longer of 2 hairs on base of ANT VI less than half as long as
base of ANT VI (Fig. 24C) . . . . . . . . . . . . . . . . . . . . . . . . . . testudinaceus
Hind tibia uniformly pale or dusky. Longer of 2 hairs on base of ANT
VI more than half as long as base of ANT VI ................... 3
3 SIPH of aptera pale. Abdomen of alata with dark transverse bars on all
tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aceris
52 Acer

SIPH of aptera with at least distal half dark. Abdomen of alata with only
faint dorsal markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . coracinus

A. pseudoplatanus Drepanosiphum acerinum, dixoni,

(European Sycamore) oregonensis, platanoidis
Periphyllus acericola, americanus,
[coracinus], [nevskyi], singeri,
testudinaceus

Key to Periphyllus species on Acer pseudoplatanus

1 Longer of 2 hairs on base of ANT VI less than half as long as base of
ANT VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Longer of 2 hairs on base of ANT VI more than half as long as base of
ANT VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Aptera with only very pale dorsal abdominal sclerites. Alata with normally
at least 8 long hairs on each of ABD TERG 1 to 7. Tibiae pale or rather
evenly pigmented, or darkening distally . . . . . . . . . . . . . . . . . americanus
Aptera with a clear pattern of dark dorsal abdominal sclerites. Alata
normally with only 6 long hairs on each of ABD TERG 1 to 7. Tibiae
with very pale middle region contrasting with dark base and distal section
................................................... testudinaceus
3 SIPH of aptera dusky to dark, shorter than their basal diameters. ANT
PT/BASE more than 4.0 (except in early spring apterae) ..... singeri
SIPH of aptera pale, a little longer than their basal diameters. ANT
PT/BASE less than 3.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . acericola

A. pseudosieboldianum Periphyllus aceris, californiensis

(Korean Maple) Yamatocallis hirayamae
(For Periphyllus spp. see key, p. 43)
A. purpurescens Yamatocallis hirayamae
A. regelii Periphyllus mamontovae, nevskyi

Key to Periphyllus species on Acer regelii

- Apterae without dorsal abdominal markings. Alatae with only 3-5
rhinaria on ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . mamontovae
- Apterae with a dark dorsal abdominal patch. Alatae with 12-13 rhinaria
on ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nevskyi

A. rubescens Trichaitophorus aceris

A. rubrum Drepanaphis acerifoliae,
(Red, Soft or Swamp Maple) carolinensis, knowltoni,
 Acer 53

nigricans, parva, saccharini,

tissoti
Periphyllus lyropictus, testudinaceus
(For Periphyllus spp. use couplet 1 of key, p. 56)

Key to Drepanaphis spp. on Acer rubrum

1 Fore femora pigmented along entire length, especially dorsally ..... 2
Fore femora pale, or dusky at distal ends only . . . . . . . . . . . . . . . . . . . 4
2 All wing veins distinctly dark-bordered. Tubercles on ABD TERG 1 well
developed, almost as long as those on ABD TERG 3 (Fig. 26F) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . acerifoliae
Wing veins not dark-bordered except for pterostigma and basal part of
Rs. Tubercles on ABD TERG 1 either much smaller than those on ABD
TERG 3, or undeveloped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ANT PT/BASE less than 7.0. Tubercles on ABD TERG 1 to 4 all
well developed; those on ABD TERG 1 and 2 finger-like and of about
equal length, about half the height of the broadly conical tubercles on
ABD TERG 3, which are united near their bases (Fig. 26E) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . carolinensis
- ANT PT/BASE more than 8.0. Only tubercles on ABD TERG 3 well
developed, united for about half their lengths; others undeveloped or
inconspicuous (Fig. 26A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . knowltoni
4 ANT I and II both as dark as head. Abdomen with dark ventral sclerites
............................................................. 5
- ANT II much paler than ANT I, like base of ANT III. Abdomen with
ventral sclerites rather faint, or absent . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 ANT III with 8-22 (usually 11+) rhinaria. Never more than 4 small
accessory rhinaria alongside the large primary rhinarium on ANT VI
....................................................... nigricans
- ANT III with 5-14 rhinaria (usually less than 11). ANT VI with 5-14
small accessory rhinaria alongside the main primary rhinarium . . . . . .
......................................................... tissoti
6 Wing veins diffusely bordered with fuscous, giving wings a slightly cloudy
appearance (best seen with hand lens). ANT III with 6-17 rhinaria (usually
10 or more) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . parva
- Wings clear. ANT III with 2-12 rhinaria (usually less than 10) . . . . . .
...................................................... saccharini
A. rufinerve Periphyllus montanus
(Red-vein Maple) Trichaitophorus koyaensis
A. saccharinum Drepanaphis acerifoliae, [parvus],
(Silver or White Maple) [sabrinae], saccharini
54 Acer

Paraprociphilus tessellatus
Periphyllus americanus, lyropictus,
testudinaceus

Key to Drepanaphis species on Acer saccharinum

Fore femora pigmented along entire length, especially dorsally. All wing
veins distinctly dark-bordered. Tubercles on ABD TERG 1 long and
finger-like, much longer than those on ABD TERG 2, almost as long as
those on ABD TERG 3 (Fig. 26F) . . . . . . . . . . . . . . . . . . . . . . acerifoliae
Fore femora pale. Wing veins not dark-bordered (except pterostigma and
basal part of Rs. Tubercles on ABD TERG 1 about equally developed to
those on ABD TERG 2, much shorter than those on ABD TERG 3
(Fig. 26G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . saccharini

Key to Periphyllus species on Acer saccharinum

1 CAUDA more than half as long as wide, with a slight basal constriction
(Fig. 23D). Longer of the 2 hairs on base of ANT VI very long and fine,
longer than base of ANT VI . . . . . . . . . . . . . . . . . . . . . . . . . . . lyropictus
CAUDA broadly rounded, less than half as long as its basal width
(Fig. 23A). Longer of 2 hairs on base of ANT VI less than half as long
as base of ANT VI (including rhinarium) . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Aptera with only very faint dorsal abdominal markings. Alata with
normally at least 8 long hairs on each of ABD TERG 1 to 7. Tibiae pale,
or rather evenly pigmented, or darkening distally . . . . . . . . americanus
Aptera with a clear pattern of dorsal abdominal sclerites. Alata with
normally only 6 long hairs on each of ABD TERG 1 to 7. Tibiae with
pale middle region contrasting with dark base and distal section .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . testudinaceus

A. saccharum Drepanaphis acerifoliae,

(Sugar or Hard Maple) carolinensis, choanotricha,
kanzensis, keshenae, knowltoni,
[pallida], parva, sabrinae,
simpsoni, tissoti
Longistigma caryae
Neoprociphilus aceris
Periphyllus americanus, lyropictus,
testudinaceus
Shenahweum minutum
(For Periphyllus spp. see key under A. saccharinum.)

Key to Drepanaphis species on Acer saccharum

1 Fore femora pigmented along entire lengths, especially dorsally .... 2
 Acer 55

Fore femora pale, or only dusky towards apices . . . . . . . . . . . . . . . . . 6

2 Wing veins distinctly dark-bordered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Wing veins not dark-bordered (except for pterostigma and base of Rs)
............................................................. 4
3 Tubercles on ABD TERG 1 to 4 all conspicuous; those on ABD TERG
1 and 3 long and finger-like, those on 1 almost as long as those on 3
(Fig. 26F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . acerifoliae
- Tubercles on ABD TERG 3 well developed, united for at least half of their
lengths; those on ABD TERG 1, 2 and 4 undeveloped or inconspicuous
(Fig.26H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . keshenae
4 ANT PT/BASE more than 9.0. Only the tubercles on ABD TERG 3 well
developed, united for about half their lengths; those on ABD TERG 1,
2 and 4 inconspicuous or undeveloped (Fig. 26A) . . . . . . . . . knowltoni
- ANT PT/BASE less than 8.0. Tubercles on ABD TERG 1 to 4 all
developed, although those on ABD TERG 3 are often largest . . . . . 5
5 Tubercles on ABD TERG 3 about twice as long as those on ABD TERG
2, which are of similar length to those on ABD TERG 1 and 4 (Fig. 26E).
On ANT VI, 4 small accessory sensoria alongside the main primary
rhinarium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . carolinensis
Tubercles on ABD TERG 2 almost as long as those on 3, clearly longer
than those on 1 and much longer than those on 4 (Fig. 26I). On ANT VI,
5-6 small accessory sensoria alongside the primary rhinarium . . . . . . . .
....................................................... sabrinae
6 ANT PT/BASE less than 6.0. All dorsal abdominal tubercles well
developed, with those on ABD TERG 1 largest (Fig. 26J) . . . . . . . . . . .
....................................................... simpsoni
- ANT PT/BASE more than 8.0. Dorsal abdominal tubercles variably
developed, but those on ABD TERG 3 always largest . . . . . . . . . . . . 7
7 ANT II concolorous with ANT III, paler than ANT I . . . . . . . . . . . . 8
- ANT II much darker than ANT III, as dark as ANT I . . . . . . . . . . . 9
8 Wing veins diffusely bordered, giving wings a cloudy appearance (view
with hand lens). Tubercles on ABD TERG 1 to 4 well developed, although
those on ABD TERG 1, 2 and 4 only one quarter to one half of the height
of those on ABD TERG 3 (Fig. 26K) . . . . . . . . . . . . . . . . . . . . . . . parva
Wings clear. Tubercles on ABD TERG 3 well developed, but those on
ABD TERG 1, 2 and 4 undeveloped or very small (Fig. 26L) . . . . . . . .
...................................................... kanzensis
9 Dorsal body hairs, including those on abdominal tubercles, clearly longer
than basal diameter of ANT III, and with distinctly expanded apices. ANT
III with 3-5 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . choanotricha
56 Acer

Dorsal body hairs mostly shorter than, or as short as, basal diameter of
ANT III, and with blunt or only very slightly expanded apices. ANT III
with 7-16 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tissoti

A. sinense Periphyllus testudinaceus

(Chinese Maple)
A. spicatum Drepanaphis spicata
(Mountain Maple)
A. stevenii Drepanosiphum oregonense
Periphyllus steveni
A. tataricum Periphyllus minutus, testudinaceus
Stomaphis graffii
A. tegmentosum Periphyllus tegmentosus
Trichaitophorus aenigmatosus
A. trautvetteri Drepanosiphum caucasicum,
oregonense
Periphyllus testudinaceus
A. tricidum Periphyllus californiensis
A. triflorum Periphyllus allogenes
A. tschonoskii Trichaitophorus japonicus
(Korean Maple, Bogjagi)
A. turkestanicum Periphyllus mamontovae, nevskyi
(see couplet under A. regelii)
A. ukurunduense Periphyllus kuwanaii
Yamatochaitophorus albus
A. velutinum Drepanosiphum iranicum
A. villosum Periphyllus villosi
Trichaitophorus aceris,
?recurvispinus
Yamatocallis brevicauda
Acer spp. Periphyllus bengalensis,
himalayensis, pallidus,
tokyoensis
Stomaphis aceris
Yamatocallis obscura, sauteri

Key to the more polyphagous Periphyllus species, and those on unidentified

species of Acer
1 CAUDA as long as its basal width, often with a slight constriction, so that
the apical part is a knob (Fig. 23D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
 Acer 57

- CAUDA shorter than its basal width, without a trace of a constriction

(e.g. Fig.23A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 SIPH dark in alata, pale to dusky in aptera, conical, about as long as their
basal width. ANT PT/BASE 4.5-6.0. Rhinaria on ANT III of alata
extending at least 0.6 of length of segment ............... lyropictus
- SIPH of both aptera and alata pale, almost cylindrical, more than
1.5 times as long as their basal width. ANT PT/BASE less than 4.0.
Rhinaria on ANT III of alata confined to basal half of segment .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . villosi
3 SIPH of aptera jet black, contrasting greatly with pale body and append-
ages (except possibly extremities of tarsi and base of ANT VI). BL less
than 2mm. 1st tarsal segments all with 5 hairs . . . . . . . . vandenboschi
- SIPH of aptera pale to dark brown, not clearly darker than other parts
of body. BL usually more than 2 mm. 1st tarsal segments with 5 or 7 hairs
............................................................. 4
4 CAUDA semicircular with only 5-6 hairs. Dorsal abdomen of aptera
without dark markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pallidus
- CAUDA broadly rounded (crescent-shaped) with at least 7 hairs. Dorsal
abdomen of aptera with or without dark markings . . . . . . . . . . . . . . . 5
5 ANT III of alata with 28-50 rhinaria, and ANT IV with 2-9 rhinaria
............................................................. 6
- ANT III of alata with 6-34 rhinaria, and ANT IV without any rhinaria
............................................................. 8
6 SIPH shorter than HT II. ANT PT/BASE of alata more than 3.0 ...
.................................................... formosanus
- SIPH as long as or longer than HT II. ANT PT/BASE of alata less than
2.8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 BL of aptera 1.8-2.2 mm, of alata 2.0-2.5 mm. Antennae of alata pale
basally, with 2-6 rhinaria on ANT IV . . . . . . . . . . . . . . . . . . bengalensis
- BL of alata 2.9-3.5 mm (aptera not described). Antennae of alata wholly
dark, with 5-9 rhinaria on ANT IV . . . . . . . . . . . . . . . . . . himalayensis
8 Hind femur with pale basal half and contrastingly dark distal half. Longer
of the 2 hairs on base of ANT VI 21-47 µm long, always less than
0.5 x length of base of ANT VI (Fig. 24C). ANT PT/BASE 2.5-4.2
(always more than 3.0 in alatae) . . . . . . . . . . . . . . . . . . . . . testudinaceus
- Hind femur either entirely pale or mostly dark except at base. Longer of
the 2 hairs on base of ANT VI 45-95 µm long, 0.3-1.3 x length of base
of ANT VI. ANT PT/BASE 1.6-3.7 (less than 3.0 except in alatae of
bulgaricus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
58 Actinodaphne

9 Hind tibiae wholly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . californiensis

- Hind tibiae pale at least in middle section . . . . . . . . . . . . . . . . . . . . . . 10
10 Apterae in spring with front of head pale, and rarely with any dorsal
markings. Dorsal hairs all pale . . . . . . . . . . . . . . . . . . . . . . . . . . acericola
- Apterae in spring usually with front of head dark, dusky to dark
transverse bars or paired patches on ABD TERG 1 to 6, and the long
spinal hairs, or at least those on ABD TERG 5-8, dark-pigmented ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Aptera with long spinal hairs on ABD TERG 5-8 (or 4-8) much darker
than spinal hairs on more anterior tergites. Longer of 2 hairs on ANT
BASE VI 0.8-1.3 x length of ANT BASE VI, and shorter hair
0.25-0.47 x BASE VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rhenanus
- Aptera with long spinal hairs similarly pigmented on all ABD TERG.
Longer of 2 hairs on ANT BASE VI 0.3-0.8 x length of BASE VI, and
shorter hair 0.12-0.27 x BASE VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 ABD TERG VI of aptera with longest hairs 270-480 µm . . . . . . . . . . . .
..................................................... bulgaricus
- ABD TERG VI of aptera with longest hairs 120-240 µm . . . . . . . . . . . .
...................................................... kuwanaii

ACTINODAPHNE Lauraceae

Actinodaphne lancifolia Aiceona japonica

A. pedicellata Aiceona actinodaphnis
Actinodaphne sp. Aiceona malayana

Key to aphids on Actinodaphne

1 R IV+V shorter than HT II. ANT PT/BASE 0.4 or less . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona japonica
- R IV+V longer than HT II. ANT PT/BASE 0.5 or more . . . . . . . . . 2
2 Hairs on ANT III less than 2 x basal diameter of ANT III . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona malayana
Hairs on ANT III mostly more than 3 x basal diameter of ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona actinodaphnis
 Aesculus 59

ACTINOPHLOEUS Palmae

Actinophloeus macarthurii Cerataphis variabilis

A. propinquus Cerataphis variabilis

ACTINOSTROBUS Cupressaceae

Actinostrobus pyramidalis Illinoia morrisoni

ADANSONIA Bombacaceae

Adansonia digitata Aphis gossypii

Macrosiphum euphorbiae
(Use key to polyphagous aphids, p. 532)

ADENIUM Apocynaceae

Adenium multiflorum Aphis gossypii

AESCULUS Horse Chestnuts Hippocastaneae

Few aphids are recorded from Aesculus, and only three are possibly specific
to members of this genus.

Host Plant List

Aesculus californica Neomyzus circumflexus
Periphyllus californiensis,
testudinaceus
Prociphilus americanus
60 Aesculus

A. glabra Drepanaphis monelli

Periphyllus aceris, testudinaceus
A. hippocastanum Aphis craccivora, fabae,
(Horse Chestnut) taraxacicola
A. indicus Periphyllus aesculi
A. parviflora Periphyllus aceris
A. turbinata Periphyllus californiensis,
[koelreuteriae]
Stomaphis yanonis ssp. aesculi

Key to aphids feeding on Aesculus

1 Rostrum much longer than body . . . . . . . . . Stomaphis yanonis aesculi
Rostrum much shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT PT/BASE much less than 1.0. SIPH absent . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus americanus
- ANT PT/BASE more than 1.0. SIPH present . . . . . . . . . . . . . . . . . . . 3
3 Adult viviparae all alate, with a large, bilobed, pigmented, dorsal
abdominal process . . . . . . . . . . . . . . . . . . . . . . . . . . . Drepanaphis monelli
Adult viviparae alate or apterous, if alate then without a dorsal abdominal
process ...................................................... 4
4 SIPH stump-shaped. Hairs on antennae much longer than basal diameter
of ANT I I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- SIPH tubular, hairs on antennae shorter than basal diameter of ANT III
............................................................. 8
5 Hind tibiae wholly black . . . . . . . . . . . . . . . . . Periphyllus californiensis
Hind tibiae at least partly pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 SIPH shorter than their width at base. ANT PT/BASE less than 2.5
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Periphyllus aesculus
- SIPH longer than their basal width. ANT PT/BASE more than 2.5 ... 7
7 Hind tibiae uniformly pale, or dusky only at apices . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Periphyllus aceris
Hind tibiae with pale middle region, contrasting with dark base and distal
section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Periphyllus testudinaceus
8 Lateral (marginal) tubercles present on abdominal segments 1-5 and 7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis taraxacicola
Lateral abdominal tubercles absent, or present only on abdominal
segments 1 and 7 . . . . . . . . . go to key to polyphagous aphids (p. 532)
 Albizia 61

AGANOPE Leguminosae

Aphis craccivora

AGATHIS Kauri Pine Araucariaceae

Agathis labillardieni Neophyllaphis rappardi

AILANTHUS Tree of Heaven Simaroubaceae

Ailanthus altissima (= glandulosa) Aphis craccivora, gossypii,

spiraecola
[Epipemphigus imaicus - error?]
Kaburagia ailanthi
[Kurisakia ailanthi - error?]
Macrosiphum euphorbiae

SIPH as small, inconspicuous pores. (Aphids in galls) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kaburagia ailanthi
SIPH tubular. (Aphids free-living) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

ALBIZIA Leguminosae

Albizia odoratissima Toxoptera aurantii

Albizia sp. Aphis gossypii
(Use key to polyphagous aphids, p. 532)
62 Alchornea

ALCHORNEA Euphorbiaceae

Alchornea cordifolia Aphis craccivora

ALEURITES Euphorbiaceae

Aleurites triloba Myzus persicae

ALNUS Alders Betulaceae

Fifty species of aphid in 24 genera are recorded as feeding on Alnus. Many

of these are strictly Alnus feeders, some also feed on Betula species. A single
key is provided, preceded by a list of Alnus species with their recorded aphids.
On the whole the records reflect the respective distributions of aphids and host
plants and there is little evidence of specificity to particular Alnus species;
hence the need for a single, all-embracing key.

Host Plant List

Alnus acuminata ssp. arguta Latgerina orizabaensis
A. cordata Crypturaphis grassii
Pterocallis alni
A. cremastogyne Eutrichosiphum alnifoliae
Mesocallis pteleae
Taoia chuansiensis
A. firma Glyphina schrankiana
Pterocallis (Recticallis)
alnijaponica, nigrostriata
Symydobius kabae
A. firmifolia Latgerina orizabaensis mexicana
Paraprociphilus mexicanus
A. formosana Greenidea (Trichosiphum) myricae
Pterocallis (Recticallis) pseudoalni
A. fruticosa Boernerina alni, alni insularia
 Alnus 63

Euceraphis ontakensis
Paraprociphilus baicalensis
A. glutinosa Betacallis alnicolens
(European Alder) Clethrobius comes
[Euceraphis betulijaponicae]
Glyphina betulae, schrankiana
Pterocallis alni, maculata
Stomaphis quercus
A. hirsuta Betacallis alnicolens
[Euceraphis betulae, caerulescens]
Hannabura alnicola
Paraprociphilus baicalensis
Pterocallis (Recticallis)
alnijaponicae, nigrostriata
Stomaphis alni
Symydobius alniarius
Tinocallis (Sappocallis) ulmicola
A. incana (N.B. incana of American Betulaphis quadrituberculata
authors = rugosa) Clethrobius comes
(Speckled Alder) Glyphina betulae, schrankiana
Pterocallis albidus, alni
Pterocallis (Recticallis)
alnijaponicae
A. japonica Betacallis alnicolens
Glyphina schrankiana
Hannabura alnicola
Mesocallis alnicola
Pterocallis (Recticallis)
alnijaponicae, nigrostriata
Symydobius alniarius, minutus
[Tinocallis zelkovae]
A. kamtschatica Boernerina alni insularia
Paraprociphilus baicalensis
A. mandshurica Boernerina alni insularia
A. matsumurae Betacallis alnicolens
Boernerina alni
Hannabura alnicola
Mesocallis alnicola
A. maximowiczi Boernerina alni, alni insularia
Euceraphis ontakensis
Paraprociphilus baicalensis
Pterocallis (Recticallis)
alnijaponicae
64 Alnus

A. nepalensis Eutrichosiphum alnicola,

[alnifoliae], raychaudhuri
Mesocallis alnicola, obtusirostris
Mollitrichosiphum acutihirsutum,
alni, alnifoliae, nandii
Neobetulaphis chaetosiphon,
pusilla
Taoia indica
A. nitida Mollitrichosiphum alni
A. orientalis Pterocallis alni
A. rhombifolia Euceraphis gillettei
(California or White Alder) Oestlundiella flava
Pterocallis alni, rhombifoliae
A. rubra Euceraphis gillettei
(Oregon or Red Alder) Oestlundiella flava
Paraprociphilus tesselatus
Pterocallis alni
A. rugosa Boernerina variabilis
Calaphis alni, alnosa
Euceraphis gillettei
Illinoia alni, wilhelminae
Paraprociphilus tessellatus
Pterocallis alni, alnifoliae,
rhombifoliae
A. serrulata Pterocallis alnifoliae
A. sibirica Betacallis alnicolens
Hannabura alnicola
Mesocallis alnicola
Pterocallis (Recticallis)
alnijaponicae
Symydobius aliarius, kabae
A. sieboldiana Pterocallis (Recticallis) sp. near
alnijaponicae
A. sinuata Boernerina occidentalis, variabilis
(Wavy-leafed or Sitka Alder)
A. subcordata Pterocallis alni
A. tenuifolia Euceraphis gillettei
(Thin-leaf Alder) Oestlundiella flava
A. tinctoria Pterocallis (Recticallis)
alnijaponicae, nigrostriata
 Alnus 65

A. viridis Boernerina depressa

(Green Alder) Paraprociphilus baicalensis

Key to Alnus-feeding aphids

1 SIPH much longer than wide (tubular) and densely covered with long hairs
............................................................. 2
- SIPH if long then without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Hind tibiae with a series of transverse ridges (Fig. 27A) . . . . . . . . . . . 3
- Hind tibiae without transverse ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 SIPH of aptera dark, curved outward, 0.4-0.5 x BL. (In alata 0.6-0.75 X
BL) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum nandii
- SIPH of aptera pale, almost straight, 0.5-0.85 x BL. (In alata
0.8-0.9 x BL) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum alni
4 SIPH of aptera with a zone of pale reticulation at base. CAUDA with
a distinct median papilla . . . . . . . . . Greenidea (Trichosiphum) myricae
- SIPH of aptera without any pale reticulation. CAUDA without a median
papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum spp.
5 Alatae with lateral balloon-like projections of the prothorax, and a pair
of tubercular frontal processes projecting forward between antennae, as
long as or longer than ANT I (Fig. 27B). Apterous morphs and immatures
with large plate-like (laminar) frontal and lateral processes, between
much-reduced antennae (Fig. 27C) . . . . . . . . . . . . . . Crypturaphis grassii
- Alata without lateral projections of prothorax, and if with frontal
processes then they are much shorter than ANT I, and if aptera has frontal
projections then they are not laminar, and the antennae are normal
............................................................. 6
6 PT very short, peg-like, not more than twice as long as its basal width;
ANT PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- PT finger-like or attenuate, if short then more than twice as long as its
basal width; ANT PT/BASE 0.5 or more . . . . . . . . . . . . . . . . . . . . . . 15
7 Rostrum much longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Rostrum much shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 HT II a little longer than base of ANT VI, and almost as long as
R IV+V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis alni
- HT II usually a little shorter than base of ANT VI, and only a little more
than half as long as R IV+V . . . . . . . . . . . . . . . . . . . Stomaphis quercus
9 Body of adult aptera more than 2.5 mm long. Wax pore plates well
developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
66 Alnus

Fig. 27. A, proximal part of hind tibia of aptera of Mollitrichosiphon montanum with stridulatory (?)
ridges; B, front of head of alata of Crypturaphis grassii; C, front of head (right side) of aptera of
C. grassii; 0, ornamentation of abdominal tergites of Glyphina schrankiana; E, the same for G. betulae;
f, head of aptera of Latgerina orizabaensis; G, head of alata of L. orizabaensis.

- Body of adult aptera less than 2.5 mm long. Wax pore plates absent ... 12
10 Antennae, legs, anal plate and subgenital plate of aptera all lightly
pigmented, and wax pore plates pale ..... Paraprociphilus baicalensis
- Antennae, legs, anal plate and subgenital plate of aptera brown to black,
and wax pore plates pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 R IV+V of aptera clearly longer than HT II, R IV bearing 4-10 accessory
hairs. Alate sexupara with narrow, transverse secondary rhinaria on
antenna, including 1-5 on base of ANT VI . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraprociphilus tessellatus
- R IV+V of aptera a little shorter than HT II, R IV bearing 2-3 accessory
hairs. Alate sexupara with secondary rhinaria on antennae more oval,
absent from base of ANT VI . . . . . . . . . . . . Paraprociphilus mexicanus
12 Eyes of aptera multifaceted. Dorsum of aptera pale with long capitate
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis albidus
 Alnus 67

- Eyes of aptera 3-faceted. Dorsum of aptera pigmented, sclerotic, with

either short spine-like hairs or long fine hairs . . . . . . . . . . . . . . . . . . . 13
13 Dorsal and lateral body hairs long and fine . . . . . . Glyphina longiseta
- Dorsal and lateral body hairs thick, spine-like . . . . . . . . . . . . . . . . . . 14
14 Dorsum of aptera adorned with short, transverse wrinkles or imbrications,
sometimes extending into a reticulation (Fig. 27D) .................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphina schrankiana
- Dorsum of aptera with a closed wart-like cuticular ornamentation
(Fig.27E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphina betulae
15 Populations containing apterous adults with branched, hair-bearing
processes projecting from sides of body and front of head (Fig. 27F).
Alatae without such processes but with an inward-facing subapical
protruberance on ANT I (Fig. 27G) . . . . . . . . . . Latgerina orizabaensis
- If populations contain apterous adults, then any processes these apterae
have are not branched, and alatae do not have an inward-facing protru-
berance on ANT I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 ANT PT/BASE less than or about 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . 17
- ANT PT/BASE at least 1.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
17 CAUDA of adult with a clearly defined constriction, delimiting the
terminal part as a knob . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
- CAUDA of adult broadly rounded or bluntly triangular, without a trace
of a constriction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
18 Anal plate bilobed, or with a medial cleft. Populations with apterae
and/or alatae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
- Anal plate entire, posterior margin rounded. Adults all alatae .... 42
19 Adult viviparae all alate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
- Populations comprising apterae only, or both apterous and alate morphs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
20 Abdomen with unpaired median dorsal processes on anterior tergites
(Figs 28A-B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- Abdomen without medial dorsal processes . . . . . . . . . . . . . . . . . . . . . . 25
21 Abdomen with median dorsal processes on ABD TERG 1-2 only, only
those on ABD TERG 2 being well developed (Fig. 28A) . . . . . . . . . . 22
- Abdomen with well-developed, finger-like median dorsal processes on
ABD TERG 1-4 or 1-5 (Fig. 28B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22 ANT III with 2-8 secondary rhinaria confined to middle part, which is
slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis alnifoliae
68 Alnus

Fig. 28. A, median dorsal process of ABD TERG 2 of Pterocallis alnifoliae; B, dorsal abdomen of
Recticallis nigrostriata; C, tip of forewing of R. nigrostriata.

- ANT III with 4-9 secondary rhinaria extending over dusky apical half of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis rhombifoliae
23 SIPH distally black, pale only at their bases . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Recticallis) pseudoalni
- SIPH entirely pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Forewings with dark markings at tips between wing veins (Fig. 28C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Recticallis) nigrostriata
- Forewings without dark markings between wing veins . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Recticallis) alnijaponicae
25 Antennae longer than body . . . . . . . . . . . . . . . . . . . . . Oestlundiella flava
- Antennae shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Antennal flagellum (ANT III-IV) wholly dark except for sensoriated basal
part of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . Boernerina occidentalis
- At least the basal parts of ANT IV and V pale . . . . . . . . . . . . . . . . . 23
27 R IV+V longer than HT II. Front of head usually pigmented. Anal vein
of forewing dark-bordered . . . . . . . . . . . . . . . . . . . . . . Mesocallis pteleae
- R IV+V shorter than HT II. Front of head pale. Anal vein of forewing
pale or dark-bordered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 ANT PT/BASE about equal to or more than 1.0. Anal vein of forewing
pale. R IV+V 0.5-0.7 x HT II . . . . . . . . . . . . . Mesocallis obtusirostris
 Alnus 69

- ANT PT/BASE only 0.6-0.7. Anal vein of forewing usually dark-

bordered. R IV+V 0.7-0.9 x HT II . . . . . . . . . . . . . Mesocallis alnicola
29 Antennae much shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
- Antennae longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
30 ABD TERG 3 of aptera with thick, apically-expanded spinopleural
hairs, 20-25 µm long, arising from tuberculate bases. Alata with a black
quadrate patch on ABD TERG 4-6, and ANT III bearing 20-25 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobetulaphis pusilla
- ABD TERG 3 of aptera with minute spinopleural hairs, 8-10 µm long.
Alata without a black dorsal abdominal patch, and with ANT III bearing
16-18 secondary rhinaria . . . . . . . . . . . . . . . Neobetulaphis chaetosiphon
31 Each SIPH with a long hair arising from it posteriorly . . . . . . . . . . 32
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
32 Knob of CAUDA elongate, about twice as long as wide. ANT VI (includ-
ing PT) less than half as long as ANT III . . . . . . . Taoia chuansiensis
- Knob of CAUDA only a little longer than wide. ANT VI (including PT)
almost as long as ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . Taoia indica
33 Aptera with ABD TERG 8 broad and sclerotic, extending posteriorly like
a canopy over CAUDA (Fig. 29A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
- Aptera with ABD TERG 8 not extending posteriorly beyond base of
CAUDA in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
34 Aptera with prominent paired frontal processes projecting forward
between antennae, square-sided in dorsal view and with a thick hair arising
at each corner (Fig.29B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
- Aptera with paired frontal processes absent, or only present as flat
tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boernerina occidentalis
35 ABD TERG 8 of aptera with 4-7 (usually 5-6) thick capitate hairs on its
posterior margin. Alata with a well-developed pair of frontal tubercles,
longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boernerina depressa
- ABD TERG 8 of aptera with 8-10 thick capitate hairs on posterior
margin. Alata with frontal tubercles undeveloped or shorter than their
basal widths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 R IV+V short, only a little longer than its basal width and less than 0.9
of length of HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boernerina alni
- R IV+V about 1.5 times longer than its basal width and about equal in
length to HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Boernerina variabilis
37 Aptera with all hairs on ANT III shorter than basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
70 Alnus

Fig. 29. A, end of abdomen of aptera of Boernerina alni; B, front of head of same; C, groups of
marginal tubercles on ABD TERG 4 and 5 of Symydobius kabae; D, dorsal abdomen of Calaphis alni.

- Aptera with at least some of hairs on ANT III longer than basal diameter
of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
38 Anterior abdominal segments of aptera each with only one capitate
marginal hair on each side . . . . . . . . . . . . . . . . . . . . Mesocallis alnicola*
- Anterior abdominal segments of aptera each with 2 capitate marginal
hairs, arising from a well-developed marginal tubercle . . . . . . . . . . . 39
39 ANT III of alata with 2-8 rhinaria confined to middle part, which is
slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis alnifoliae
- ANT III of alata with 4-9 rhinaria extending over dusky apical half
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis rhombifoliae
 Alnus 71

40 Aptera with apices of antennal segments and SIPH unpigmented, and

ANT PT/BASE 0.2-0.3. Alata with 6-9 secondary rhinaria . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis albidus
- Aptera with apices of antennal segments and SIPH dark, and ANT
PT/BASE 0.5-0.8. Alata with 2-5 secondary rhinaria . . . . . . . . . . . . 41
41 Aptera with dorsal body hairs pigmented. ANT III with more than 2 long
hairs, and some hairs on ANT IV and V longer than basal diameters of
their respective segments . . . . . . . . . . . . . . . . . . . . . . Pterocallis maculata
- Aptera with dorsal body hairs pale. ANT III with only 1-2 long hairs,
and all hairs on ANT IV and V short and inconspicuous . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis alni
42 Hairs on ANT III much longer than basal diameter of segment . . . . .
.............................................. Clethrobius comes
- Hairs on ANT III much shorter than basal diameter of segment . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
43 Thorax and legs dark . . . . . . . . . . . . . . . . . . . . . . . Euceraphis ontakensis
- Thorax and legs pale . . . . . . . . . . . . . . . . . . . . . . . . . . Euceraphis gillettei
44 Small aphid, less than 2mm long. Abdomen without dorsal transverse
bands. Anal plate bilobed . . . . . . . . . . . . . Betulaphis quadrituberculata
- Large aphid, more than 2 mm long. Abdomen with dark dorsal transverse
bands. Anal plate rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
45 Abdominal marginal sclerites each bearing 1-7 small transparent tubercles
(Fig. 29C). ANT III with 16-35 rhinaria . . . . . . . . . Symydobius kabae
- Abdominal marginal sclerites without tubercles. ANT III with 18-90
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
46 Hind tibia mainly pale. ANT III with 18- 40 rhinaria. ANT VI BASE
with 3-9 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symydobius minutus
- Hind tibia wholly blackish-brown. ANT III with 30-90 rhinaria. ANT VI
BASE with 15-24 hairs . . . . . . . . . . . . . . . . . . . . . . Symydobius alniarius
47 SIPH short, truncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
- SIPH long, tubular, and usually slightly to markedly swollen on apical
third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
48 Head with a transverse black band ventrally between eyes. ANT
PT/BASE less than 2.0 . . . . . . . . . . . . . . . . . . . . . . . Betacallis alnicolens
- Head uniformly pale or dark, without a ventral transverse black band.
ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
49 Abdomen with an extensive black dorsal patch, including the SIPH
(Fig. 29D) ......................................... Calaphis alni
72 Alsophila

- Abdomen without a black patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

50 Antennae dark. Aptera with 4-6 dorsal capitate hairs per segment ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis alnosa
- Antennae pale. Aptera with 12-30 dorsal capitate hairs per segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hannabura alnicola
51 Antennae dusky. R IV+V 1.1-1.3 x HT II, with 7-10 accessory hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 . . . Illinoia alni
- Antennae pale (except for extreme apices of segments). R IV+V
c. 2.0 x HT II, very hairy, with 30 or more accessory hairs . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia wilhelminae

ALSOPHILA = CYATHEA

AMELANCHIER June Berries Rosaceae

Host Plant List

Amelanchier alnifolia Acyrthosiphon macrosiphum
(Saskatoon Berry) Aphis whiteshellensis
Nearctaphis sensoriata
Prociphilus caryae ssp. fitchi
A. florida Acyrthosiphon macrosiphum
(Serviceberry) Fimbriaphis gentneri
A. laevis Acyrthosiphon macrosiphum
(Serviceberry) Aulacorthum solani
Eriosoma americanum
Fimbriaphis gentneri
Prociphilus caryae
A. ovalis Dysaphis parasorbi
Fimbriaphis gentneri
A. spicata Macrosiphum amelanchiericolens
Nearctaphis sensoriata

Key to the aphids feeding on Amelanchier species

1 ANT PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
 Amygdalus 73

2 SIPH present as raised pores with rims partly sclerotized, ringed by hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma americanum
SIPH absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus caryae
3 CAUDA triangular, helmet-shaped or rounded, shorter than or about as
long as its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
CAUDA at least 1.5 times longer than its basal width . . . . . . . . . . . . 6
4 SIPH 2-3 times longer than CAUDA . . . . . . . . . . . Dysaphis parasorbi
SIPH about as long as, or a little longer than, CAUDA . . . . . . . . . . 5
5 Dorsum with extensive dark pigmentation. Basal part of hind tibia slightly
swollen, bearing a group of circular 'pseudosensoria' (scent glands)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis sensoriata
Dorsum with pigmented areas restricted to posterior abdominal segments.
Hind tibiae of viviparous morphs without scent glands . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis whiteshellensis
6 SIPH and CAUDA both black. Antennal tubercles weakly developed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
SIPH pale or dark, but if dark then the CAUDA is paler. Antennal
tubercles well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ANT III in aptera with about 40 rhinaria, distributed over entire length
of segment. SIPH with an apical zone of polygonal reticulation
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum amelanchiericolens
- ANT III in aptera with 1-2 rhinaria near base . . . . . . . . . . . . . . . . . . . 8
8 SIPH of aptera dark and very long, nearly half as long as body . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon macrosiphum
SIPH of aptera about one quarter of body length, pale except sometimes
at their apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Aptera with antennal tubercles diverging in dorsal view (Fig. 42K). SIPH
wholly pale. Alata with a solid dark dorsal abdominal patch . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri
Aptera with antennal tubercles parallel in dorsal view (Fig. 42J). SIPH
often dark-tipped. Alata with dark transverse bars on dorsal abdomen,
sometimes fused across segments in midline only . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani

AMYGDALUS Almonds Rosaceae

See Prunus dulcis

74 Anacardium

ANACARDIUM Anacardiaceae

Anacardium occidentale Aphis gossypii, spiraecola

Brachyunguis harmalae
Toxoptera aurantii, odinae

- ANT PT/BASE a little less than 1.0 . . . . . . . . Brachyunguis harmalae

- ANT PT/BASE much more than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

ANDIRA Leguminosae

Andira inermis Aphis craccivora

ANNONA Custard Apples Annonaceae

Key to aphids on Annona (This is a revision of the key in B & E, 1984)

1 SIPH with numerous long hairs . . . . . . . . . . . . . . . . Greenidea annonae
- SIPH without hairs ........................................... 2
2 SIPH in form of small, broad-based cones. CAUDA knobbed, anal plate
bilobed. Aptera with posterior abdomen bearing backwardly directed
dorsal processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis kalipadi
SIPH tubular, tapering. CAUDA tongue-like, anal plate entire. Aptera
without posterior abdominal processes . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Aptera with an extensive black dorsal abdominal patch (Fig. 121N)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
Aptera without a black dorsal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 CAUDA black, like siphunculi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
CAUDA pale, or if dusky then paler than siphunculi . . . . . . . . . . . . . 6
5 ANT PT/BASE less than 3.0. Stridulatory apparatus absent . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
 Apocynum 75

- ANT PT/BASE more than 3.5. Stridulatory apparatus present (Fig. 122)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
6 Lateral (marginal) tubercles present on most abdominal segments ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis sassceri
- Lateral tubercles usually only on ABD TERG 1 and 7 . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii

ANOGEISSUS Combretaceae

Anogeissus schimperi Aphis gossypii

ANTIDESMA Euphorbiaceae

Antidesma bunius Sinomegoura citricola

APHANANTHE Ulmaceae

Aphananthe aspera Stomaphis aphananthae

APOCYNUM Apocynaceae

Aphis asclepiadis, fabae, nerii,

spiraecola
Aulacorthum solani
Paulianaphis madagascariensis
Macrosiphum euphorbiae
Myzus ornatus, persicae
Rhopalosiphoninus staphyleae
Toxoptera aurantii
76 Aralia

Key to aphids on Apocynum

1 Dorsal abdomen with 4 very long, spinulose, spine-like processes posterior
to the siphunculi . . . . . . . . . . . . . . . . . . . . Paulianaphis madagascariensis
Abdomen without such processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Alata with more than 20 rhinaria on ANT III, not in a single row ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ...... Aphis asclepiadis
- Alata with less than 20 rhinaria on ANT III, arranged in approximately
single file . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

ARALIA Araliaceae

Aralia elata Aphis fabae

Aulacorthum solani
Cavariella araliae
Myzus persicae
Rhopalosiphoninus staphyleae
Toxoptera odinae

1 Supracaudal process present, that of aptera exceeding length of CAUDA

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cavariella araliae
No supracaudal process .... go to key to polyphagous aphids, p. 532
(but specimens coming to Aphis fabae in this key may be A. hederae
(q.v.), although this species has not yet been recorded from Aralia)

ARAUCARIA Monkey Puzzles etc. Araucariaceae

Host Plant List

Araucaria angustifolia No aphids recorded
(Parana Pine)
A. bidwilli Neophyllapis araucariae
A. cunninghamii Neophyllaphis araucariae
A. excelsa Neophyllaphis araucariae
A. heterophylla Illinoia morrisoni
Neophyllaphis araucariae
 Arenga 77

Key to aphids on Araucaria

Eyes of aptera 3-faceted. SIPH pore-like. Antennae usually 5-segmented
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neophyllaphis araucariae
Eyes of all morphs multifaceted. SIPH long and tubular, swollen distally.
Antennae 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni

ARBUTUS Ericaceae

Arbutus andrachne Wahlgreniella nervata (ssp. arbuti)

A. menziesi Aphis madronae
Wahlgreniella nervata (ssp. arbuti)
A. unedo Aphis arbuti
(Strawberry Tree) Wahlgreniella nervata (ssp. arbuti)

Key to aphids on Arbutus species

1 CAUDA triangular or helmet-shaped, shorter than its basal width in
dorsal view, and bearing numerous hairs . . . . . . . . . . . Aphis madronae
CAUDA much longer than its basal width . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH dark and tapering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis arbuti
SIPH pale with dark apices, slightly swollen on distal half . . . . . . . . . .
............................................ Wahgreniella nervata

ARECA Palmae

Areca catechu Cerataphis variabilis

(For aphids on palms see B & E, 1984)

ARENGA Palmae

Arenga pinnata Cerataphis variabilis

(For aphids on palms see B & E, 1984)
78 Aristotelia

ARISTOTELIA Elaeocarpaceae

Aristotelia serrata Paradoxaphis aristoteliae

ARTOCARPUS Breadfruit Moraceae

Artocarpus spp. Greenidea artocarpi

Toxoptera aurantii

Key to species
SIPH with numerous long hairs. Stridulatory apparatus absent . . . . . .
............................................. Greenidea artocarpi
SIPH without hairs. Stridulatory apparatus present (Fig. 122) . . . . . . . .
.............................................. Toxoptera aurantii

ARUNDINARIA Bamboos Gramineae

Arundinaria is here used in its older, broad sense, including Pleioblastus and
some other groups now distinguished as separate genera, e.g. Yushania. It
seems to have a more restricted aphid fauna than Bambusa and some species
are specific to it, so a separate, shorter key is provided. However, it is probable
that other bamboo-feeding aphids, not so far recorded from Arundinaria, will
feed on this genus and it may be worthwhile to work through the Bambusa
key if your aphid is not one of those listed and keyed here.

The following aphids are recorded from Arundinaria (and related genera)
Astegopteryx minuta, pallida
Cerataphis bambusifoliae
Ceratovacuna sylvestrii
Chaitoregma tattakana
Cranaphis formosana, [indica]
Melanaphis arundinariae,
bambusae, meghalayensis ssp.
bengalensis, ?pahanensis
 Arundinaria 79

Neocranaphis arundinariae
Paracolopha morrisoni
Pseudoregma koshuensis
Rhopalosiphum arundinariae
Sitobion papillatum ssp.
subnudum
Takecallis affinis, arundicolens,
arundinariae, sasae, taiwanus

Key to species (species not seen by authors marked with an *)

1 Antenna of aptera more than 0.5 x BL, or if only alatae are present, these
have antennae bearing circular or oval secondary rhinaria . . . . . . . . . 2
- Antennae of aptera much less than 0.5 X BL, and secondary rhinaria on
the antennae of alatae are ring-like, extending more than half-way around
the circumference of the antenna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2 Anal plate bilobed, CAUDA knobbed. ANT PT/BASE 1.0 or less. (All
viviparae alate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Anal plate entire, CAUDA tongue-shaped (sometimes with a mid-way
constriction). ANT PT/BASE at least 1.5 . . . . . . . . . . . . . . . . . . . . . . . 9
3 ANT PT/BASE only about 0.25 . . . . . . . Neocranaphis arundinariae*
- ANT PT/BASE at least 0.75 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Antennae and tibiae wholly dusky to dark, and SIPH dusky . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cranaphis formosanus*
- Antennae pale to dusky or contrastingly banded, tibiae pale to dusky,
SIPH pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Antennae longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Antennae shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
6 Antennae III-VI dark except for basal part of ANT III; ANT III with
10-15 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Takecallis affinis*
- Antennae III-VI or IV-VI pale basally and dark apically; ANT III with
3-9 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Thorax with longitudinal dark stripes, and abdomen with a pair of
elongate dark patches on each tergite. ANT III wholly dusky, CAUDA
pale or dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Takecallis arundinariae
- Thorax and abdomen without dark dorsal markings. ANT III with a black
sensoriated section contrasting with pale base and apical half. CAUDA
black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Takecallis arundicolens
8 Secondary rhinaria confined to basal third of ANT III. Only 2 spinal hairs
on each abdominal tergite . . . . . . . . . . . . . . . . . . . . . Takecallis taiwanus
80 Arundinaria

- Secondary rhinaria extending about half of length of ANT III. Each

abdominal tergite bearing at least 4 hairs besides the marginal ones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Takecallis sasae
9 SIPH similar in length to, or shorter than, CAUDA . . . . . . . . . . . . . 10
- SIPH about 2 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10 Hairs on ANT III short and inconspicuous, maximally about 0.5 x basal
diameter of segment. Alata with dark wing veins . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis bambusae
- Hairs on ANT III at least 2-3 times longer than basal diameter of segment.
Alata with wing veins not deeply pigmented . . . . . . . . . . . . . . . . . . . . 11
11 Antennae 5-segmented. SIPH more than 1.5 x longer than their width at
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis arundinariae
- Antennae 6-segmented. SIPH less than 1.5 x longer than their width at
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 SIPH a little longer than their width at base . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis pahanensis
- SIPH a little shorter than their width at base . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis meghalayensis
13 Body spindle-shaped, CAUDA much paler than siphunculi and about
3 x its basal width in dorsal view. Aptera with a single rhinarium on
ANT III . . . . . . . . . . . . . . . . . . . . . . Sitobion papillatum ssp. subnudum
- Body small and oval, CAUDA as dark as siphunculi and only a little
longer than its basal width. Aptera without any rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum arundinariae
14 Antennae of aptera extremely short, hardly longer than the elongate R
IV+V. Tarsi 1-segmented. SIPH displaced towards the midline (on roots)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracolopha morrisoni
- Antennae of aptera short, but much longer than the short R IV+V. Tarsi
2-segmented. SIPH in more normal position . . . . . . . . . . . . . . . . . . . . 15
15 Dorsal cuticle of head, thorax and abdomen of aptera forming an almost
circular, flattened, sclerotic shield, with only ABD TERG 8 free, and with
a distinctly crenulate margin due to a continuous row of wax glands
(Fig. 30A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerataphis bambusifoliae
- Body oval, with at least meso- and metathoracic and anterior abdominal
tergites separate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Dorsal cuticle uniformly sclerotized, with numerous separate wax glands
scattered over entire surface. Head with rounded frontal horns, even in
young immatures . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitoregma tattakana
- At least abdominal tergites mainly membranous, with localized groups of
 Arundinaria 81

Fig. 30. A, dorsal (right) and ventral (left) views of aptera of Cerataphis bambusifoliae; B, marginal
wax pore plates on right side of abdomen in Astegopteryx spp., from left to right minuta, pallida,
bambusae, bambucifoliae; C, front of head of aptera of Pseudoregma koshuensis; D, front of head of
aptera of Ceratovacuna sylvestrii; E, marginal tubercles of Chucallis bambusicola; F, wax gland on ABD
TERG 8 of Pseudoregma pendleburyi; G, the same in Ceratovacuna japonica.

wax pore plates. Frontal horns conical, often pointed in adults and always
so in young immatures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 SIPH pores placed on shallow cones encircled with fine hairs. Aptera with
wax pore plates (if present) marginal except on posterior abdominal
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
82 Bambusa

- SIPH cones small and sclerotic, not encircled by hairs. Aptera with spinal
as well as marginal wax pore plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18 Apices of frontal horns of adult aptera rounded. Marginal wax pore plates
on anterior abdominal segments either small, oval and discontiguous
(Fig. 3OB), or absent . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx minuta
- Apices of frontal horns of aptera pointed. Marginal wax pore plates on
anterior abdominal segments large, mainly oblong, closely opposed to one
another, in rows of 3-7 per segment (Fig. 3OB) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx pallida
19 Dorsal cuticle of fused head and prothorax with denticular ornamenta-
tion. Frontal horns short, curved outwards and narrowed apically, about
as long as ANT I (Fig. 30C) . . . . . . . . . . . . . . Pseudoregma koshuensis
- Dorsal cuticle of fused head and prothorax smooth or slightly wrinkled,
without denticular ornamentation. Frontal horns longer, erect, usually
exceeding length of ANT I (Fig. 3OD) . . . . . . . . Ceratovacuna sylvestrii

BAMBUSA Bamboos Gramineae

About 40 species of aphid colonize members of the genus Bambusa. Many of

these also occur on other bamboo genera but the large number recorded from
Bambusa may simply be because it is the best-known genus. The key is an
updated and expanded version of that provided by Blackman and Eastop
(1984). It includes species recorded from unidentified bamboos, possibly not
of this genus. The Hormaphidinae associated with bamboos are in need of
further study to determine their life cycles and clarify taxonomic relationships,
and reliable identification of Astegopteryx, Ceratovacuna and Pseudoregma
to species is not usually possible in the present state of knowledge.

The following aphids are recorded from Bambusa spp.

Drepanosiphinae Chucallis bambusicola
Neocranaphis bambusicola
Phyllaphoides bambusicola
Subtakecallis brevisetosa, pilosa
Takecallis affinis, arundicolens,
arundinariae, sasae, taiwanus
Aphidinae Hysteroneura setariae
Melanaphis bambusae
Rhopalosiphum rufiabdominalis
Sitobion bambusicola, [miscanthi]
Hormaphidinae Aleurodaphis antennata
 Bambusa 83

Astegopteryx bambusae,
bambucifoliae, basalis, flava,
glandulosa, [insularis],
liukueinensis, [malaccensis],
minuta, [neelagiriensis], pallida,
[salatigensis], [similis],
singaporensis, [striata],
unimaculata, [vandermeermohri]
Cerataphis bambusifoliae
Ceratoglyphina bambusae,
bengalensis, [styracicola]
Ceratovacuna [brevicornis],
floccifera, hoffmanni, indica,
japonica, keduensis, [longifila],
sylvestrii
Chaitoregma [aderuensis], tattakana
Glyphinaphis bambusae
[Indoregma bambusae]
[Pseudoastegopteryx himalayensis],
Pseudoregma albostriata,
bambusicola, [cantonensis],
gombakana, koshuensis,
montana, pendleburyi, [uscare]
Pemphiginae (Fordini) Forda hirsuta
Smynthurodes betae

Key to aphids on Bambusa (species not seen by authors marked with an *)

1 Aptera with multifaceted eyes and antennae more than 0.4 x BL or, if
only alatae are present, these have usually 6-segmented antennae bearing
circular, oval or elongate oval secondary rhinaria . . . . . . . . . . . . . . . . 2
- Apterae with 3-faceted eyes and antennae less than 0.4 x BL. Alatae
usually with 5-segmented antennae bearing ring-like rhinaria which extend
more than half-way around the circumference of the antenna (exceptions
are Fordini on roots) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2 Anal plate bilobed, CAUDA knobbed. ANT PT/BASE less than 1.0. (All
viviparae alate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Anal plate entire, CAUDA tongue-shaped (sometimes with a mid-way
constriction). ANT PT/BASE at least 1.5 . . . . . . . . . . . . . . . . . . . . . . . 8
3 Abdomen with long, finger-like, hair-bearing spinal and marginal
tubercles, especially large marginally on ABD TERG 4 (Fig. 30E) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chucallis bambusicola
- Abdomen at most with only small tubercles, none longer than width at
base ......................................................... 4
84 Bambusa

4 ANT PT/BASE 0.6 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

- ANT PT/BASE at least 0.75 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5 Clypeus without an anterioventral projection. ANT I more than
1.5 x longer than wide . . . . . . . . . . . . . . . . . Neocranaphis bambusicola
- Clypeus with a nose-like anterioventral projection. ANT I about as long
as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Dorsal abdominal hairs 30-35 µm long, at most equal to basal diameter
of ANT III. ANT PT/BASE 0.5-0.6. Dorsal abdominal sclerites small
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subtakecallis brevisetosus
- Dorsal abdominal hairs 70-95 µm, much longer than basal diameter of
ANT III. ANT PT/BASE 0.24-0.30. Dorsal sclerites large, at least on
ABD TERG 3-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subtakecallis pilosa
7 Eyes without ocular tubercles. SIPH cones very flat, mammariform
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaphoides bambusicola
- Eyes with ocular tubercles. SIPH cones truncate . . . . . Takecallis spp.
(See couplets 5-8 of Arundinaria key)
8 Antennae of aptera 5-segmented with longest hairs 3 or more X basal
diameter of ANT III. ANT PT markedly curved . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum rufiabdominalis
- Antennae of aptera 6-segmented with much shorter hairs. ANT PT not
markedly curved . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 SIPH short and truncate, similar in length to the equally dark CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis bambusae
- SIPH at least 1.5 x the much paler CAUDA . . . . . . . . . . . . . . . . . . . 10
10 Body oval. SIPH without an apical zone of polygonal reticulation.
CAUDA long and pale, with only 4 hairs ..... Hysteroneura setariae
- Body elongate oval. SIPH with apical zone of polygonal reticulation.
CAUDA dusky with about 8 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion spp. (see Sitobion bambusicola)
11 SIPH pores absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- SIPH pores present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12 ANT III less than 2 x ANT II. Dorsum densely hairy. BL 1.5-2.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smynthurodes betae
- ANT III more than 2 x ANT II. Dorsum sparsely hairy. BL 2.7-3.7 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forda hirsuta
13 Body of aptera including head with a continuous row of small wax glands
forming a complete, crenulate margin . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Body of aptera without a complete crenulate margin of wax glands ... 15
 Bambusa 85

14 Body of aptera narrowly oval; BL about 2 x greatest width. Entire dorsal

cuticle ornamented with numerous small, nodulose wax glands. R IV+V
longer than HT II . . . . . . . . . . . . . . . . . . . . . . . . Aleurodaphis antennata
- Body of aptera broadly oval to almost circular; BL maximally about
1.5 x greatest width. Dorsum without nodules. R IV+V shorter than
HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerataphis bambusifoliae
15 Head of aptera without frontal horns. Dorsal and marginal body hairs
thick and spine-like . . . . . . . . . . . . . . . . . . . . . . . Glyphinaphis bambusae
- Head of aptera with a pair of forwardly-directed frontal horns. Body hairs
short, or if long then fine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 CAUDA and anal plate rounded. Frontal horns of adult aptera usually
longer than ANT I and II combined . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- CAUDA knob-like, anal plate bilobed. Frontal horns of adult aptera not
longer than ANT I + II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 Frontal horns of aptera with 8-10 hairs, the longest 22-33 µm. SIPH with
4-6 surrounding hairs . . . . . . . . . . . . . . . . . . . Ceratoglyphina bambusae
- Frontal horns of aptera with 14-18 hairs, the longest 60-65 µm. SIPH with
6-11 surrounding hairs . . . . . . . . . . . . . . . . . Ceratoglyphina bengalensis
18 SIPH pores placed on shallow cones encircled by long fine hairs. Aptera
with wax pore plates if present usually arranged in longitudinal marginal
rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
- SIPH pores on small pigmented cones without a ring of fine hairs. Wax
pore plates of aptera if present not arranged in longitudinal marginal rows
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
19 Frontal horns of aptera with rounded apices . . . . . . . . . . . . . . . . . . . . 20
- Frontal horns of aptera with pointed apices . . . . . . . . . . . . . . . . . . . . 23
20 Marginal wax pore plates of aptera in clusters . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx liukueinensis*
- Marginal wax pore plates of aptera either in rows or completely absent
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 ABD TERG 6 of aptera usually with 3 or more hairs, and ABD TERG
2 and 3 each with 9-15 hairs (in addition to the hairs near the wax glands).
Marginal wax pore plates often touching each other over half their width,
so that the adpressed margins are straight (Fig. 30B) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx bambucifoliae/unimaculata
- ABD TERG 6 of aptera usually with 2 hairs, and ABD TERG 2 and 3
each with 4-11 hairs (in addition to those near the wax glands). Marginal
wax pore plates if developed mostly touching over much less than half
their width, and therefore more rounded . . . . . . . . . . . . . . . . . . . . . . . 22
86 Bambusa

22 Marginal wax pore plates of aptera small (10-15 µm diameter), rounded,

not touching each other (Fig. 30B), and sometimes completely absent from
some or all segments . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx minuta
- Marginal wax pore plates of aptera larger, diameter mostly 20-40 µm,
usually touching over some distance in middle (Fig. 30B), present on most
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx bambusae
23 Frontal horns of adult aptera narrowed distally and acutely pointed
apically, more than 1.5 x ANT I, and usually about as long as ANT I + II
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx pallida
- Frontal horns of adult aptera short, conical, broad-based, about equal in
length to or shorter than ANT I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Frontal horns of aptera much shorter than ANT I, only 0.04-0.08 x head
width across eyes. Marginal wax pore plates form clusters when well
developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx glandulosa
- Frontal horns of aptera similar in length to ANT I, 0.09-0.24 x head
width across eyes. Marginal wax pore plates form longitudinal rows when
well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 SIPH of aptera usually with 2-3 surrounding hairs (rarely up to 7), and
2-3 hairs on ABD TERG 5 between SIPH. Total antennal length
0.65-0.85 x width of head across eyes. R IV+V 0.73-0.92 x HT II
............................................ Astegopteryx basalis
- SIPH of aptera with 4-11 surrounding hairs, and 3-7 hairs on ABD TERG
5 between SIPH. Total antennal length 0.7-1.1 x head width across eyes.
R IV+V 0.55-0.70 x HT II . . . . . . . . . . . . . Astegopteryx singaporensis
26 Dorsal cuticle of aptera uniformly sclerotized . . . . . . . . . . . . . . . . . . . 27
- Aptera with at least the abdominal tergites partly membranous ... 28
27 Frontal horns of aptera thumb-shaped; cylindrical with broadly rounded
apices, even in immature individuals. Wax pores all dispersed, not
forming wax pore plates, even on ABD TERG 8 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitoregma tattakana
- Frontal horns of aptera conical, their apices rounded in adults, narrower
and more pointed in immatures. Wax pore plates present, at least as a
central group on ABD TERG 8 (Fig. 30F) . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoregma pendleburyi
28 Sclerotic areas of cuticle, especially head and prothorax, evenly covered
with small irregularly shaped warts or denticles . . . . . . . . . . . . . . . . . 29
- Sclerotic areas of cuticle smooth or slightly wrinkled . . . . . . . . . . . . 30
29 Aptera often without spinal wax glands. ABD TERG 1-5 may have 1-4
wax pore plates in paired spinal groups; there may be 1-2 centrally on
ABD TERG 7 and 1-4 centrally on ABD TERG 8. Alatae have secondary
 Barringtonia 87

rhinaria distributed III 25-48, IV 11-20, V 4-13, and siphunculi with a

posterior extension of the perisiphuncular sclerite . . . . . . . . . . . . . . . . . .
....................................... Pseudoregma bambusicola
- Aptera usually with well-developed wax glands on paired spinal sclerites,
only rarely without. ABD TERG 1-5 usually have 1-13 large wax pore
plates in each spinal group, and 3-4 on ABD TERG 8, but usually none
on ABD TERG 7. Alatae have secondary rhinaria distributed III 22-30,
IV 6-12, V 7-11, and siphunculi with a very small perisiphuncular sclerite
without any posterior extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoregma montana/koshuensis
30 Aptera with wax pore plates well developed only on margins of body,
except for a large medial group on the ABD TERG 8 . . . . . . . . . . . 31
- Aptera with wax pore plates well developed spinally as well as marginally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
31 Aptera with SIPH cones lightly sclerotized and bearing some long, fine
hairs. Wax pore plates on ABD TERG 8 arranged in a row . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratovacuna indica*
- Aptera with SIPH cones low, well sclerotized and without hairs. Wax pore
plates on ABD TERG 8 not in a row (e.g. Fig. 30G) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Ceratovacuna japonica/longifila/floccifera
32 Aptera with wax pore plates very large and numerous, almost occupying
the whole of the dorsal cuticle . . . . . . . . . . . . . . Ceratovacuna sylvestrii
- Aptera with fewer wax pore plates; most abdominal tergites with one
spinal group and one marginal group on each side . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratovacuna hoffmanni/keduensis

BANISTERIOPSIS Malpighiaceae

Banisteriopsis sp. Aphis gossypii

BARRINGTONIA Lecythidaceae

Barringtonia asiatica Toxoptera aurantii

88 Bauhinia

BAUHINIA Leguminosae

Bauhinia sp. Aphis gossypii, spiraecola

(Use key to polyphagous aphids, p. 532)

BERCHEMIA Rhamnaceae

Berchemia lineata Sitobion berchemiae

B. zeyheri Toxoptera aurantii, citricidus

BERSAMA Melianthaceae

Bersama ugandensis Aulacorthum solani

Sitobion halli, leonidasi

Key to aphids on Bersama

1 Inner faces of antennal tubercles parallel in dorsal view (Fig. 42J). SIPH
without apical reticulation . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
Inner faces of antennal tubercles divergent in dorsal view (Fig. 42K). SIPH
with an apical zone of polygonal reticulation .................... 2
2 Aptera with extensive black dorsal pigmentation and dark antennae, legs,
SIPH and CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion leonidasi
Aptera without extensive dark pigmentation . . . . . . . . . . Sitobion halli

BETULA Birches Betulaceae

Sixty-one species of aphids in 18 genera feed on Betula. Although the degree

of host plant specificity may be somewhat greater than in Alnus-feeding
aphids, only a small number of aphid species restrict their feeding to a single
species of Betula. Therefore, as in the case of Alnus, a single key is provided
to all birch-feeding aphids.
 Betula 89

Host Plant List

bg
indicates records from exotic hosts in botanic gardens)
Betula alba see B. pendula, B. pubescens
B. albosinensis Calaphis flavabg
(Chinese Red-Barked Birch) Euceraphis betulaebg
B. alnoides Betacallis querciphaga, sikkimensis
Betulaphis hissarica
Clethrobius dryobius
Euceraphis betulaebg
Hamamelistes sp. (miyabei?)
[Mesocallis obtusirostris]
Neobetulaphis chaetosiphon,
immaculata, pusilla
B. alpestris Hamamelistes betulinus
Hormaphis betulae
B. americana ?Hamamelistes spinosus
B. andrewsii Calaphis flavabg
Euceraphis betulaebg
B. caerulea Calaphis flavabg
Callipterinella minutissimabg
Euceraphis betulaebg
B. costata Euceraphis betulijaponicae
B. dahurica Betulaphis japonica
Calaphis betulicola, flavabg, similis
Callipterinella calliptera.
[tuberculata]
Euceraphis betulae group bg,
betulijaponicae
Glyphina betulae
Hamamelistes betulinus
Hormaphis betulae
Neobetulaphis alba
Symydobius kabae, oblongus
B. ermanii Betacallis odaiensis
Betulaphis japonica
Calaphis betulicola, flavabg,
similis
Clethrobius comes
Euceraphis ontakensis
Hamamelistes betulinus, gibberi
Symydobius kabae
90 Betula

B. forestii Calaphis flavabg

Callipterinella minutissimabg
Clethrobius comesbg
Euceraphis punctipennisbg
Hormaphis betulaebg
B. fruticosa Betulaphis japonica,
quadrituberculatabg
Symydobius oblongus
B. glandulosa Betulaphis pelei
(American Dwarf Birch) Calaphis betulaefoliae,
viridipallida
Euceraphis sp. near betulae
B, gmelini Symydobius kabae
B. grossa Calaphis flavabg
Euceraphis betulaebg
Hamamelistes gibberi ssp. grossa
B. hissarica Betulaphis hissarica
B. jaquemontii Calaphis flavabg
(White-barked Himalayan Birch) Euceraphis betulaebg
B. kamtschatica Callipterinella callipterus
Clethrobius comes
Hamamelistes betulinus
B. latifolia Calaphis flavabg
Stomaphis takahashii
B. lenta Euceraphis mucida
(Black Birch, Cherry Birch)
B. litwinowii Betulaphis quadrituberculata
B. luminifera Betacallis luminiferus
Euceraphis sp.
B. lutea (=alleghaniensis) Betulaphis quadrituberculata
(Yellow Birch) Calaphis betulaecolens, flava,
viridipallida
Euceraphis betulaebg,
betulijaponicaebg, caerulescensbg
B. mandshurica Betulaphis japonica
Calaphis betulicola, similis
Callipterinella calliptera,
[tuberculata]
Clethrobius comes
Euceraphis betulae, betulijaponicae,
caerulescens
 Betula 91

Glyphina betulae
Hamamelistes betulinus
Monaphis antennata
Symydobius kabae, oblongus
B. maximowicziana Betacallis odaiensis
Betulaphis japonica
Calaphis flavabg
Clethrobius comes
Hamamelistes betulinus ssp.
makabiae, cristafoliae
Monaphis antennata
Symydobius kabae
B. middendorfii Betulaphis quadrituberculatabg
Calaphis flavabg
Euceraphis caerulescens
Glyphina betulaebg
Monaphis antennatabg
B. nana Betulaphis pelei
(European Dwarf Birch) Calaphis arctica, betulicola, ?flava
Euceraphis sp. near betulae
Symydobius sp.
B. nigra Calaphis betulella, neobetulella
(River Birch) Callipterinella tuberculata
Hamamelistes spinosus
Hormaphis hamamelidis
Pemphigus ephemeratus
B. occidentalis (=fontinalis) Betulaphis quadrituberculata
(Western Black Birch) Calaphis betulaecolens,
betulaefoliae, coloradoensis,
flava
Euceraphis sp. near betulae
Symydobius intermedius
B. papyrifera Calaphis betulaecolens,
(Paper-bark Birch) betulaefoliae, flava, viridipallida
Callipterinella callipterus,
minutissima
Euceraphis sp. near betulae,
punctipennisbg
Glyphina setosa
Hamamelistes spinosus
Hormaphis hamamelidis
Monaphis antennata
Symydobius americana
92 Betula

B. pendula (=verrucosa) Betulaphis brevipilosa,

(Silver Birch) quadrituberculata
Calaphis betulaecolens, betulicola,
flava
Callipterinella callipterus,
minutissima, tuberculata
Clethrobius comes,
Euceraphis betulae
Glyphina betulae
Hamamelistes betulinus, spinosus
Monaphis antennata
Stomaphis quercus
Symydobius oblongus
B. platyphylla (including var. Betacallis odaiensis
japonica) Betulaphis japonica
Calaphis betulicola, flava, similis
Callipterinella callipterus,
[tuberculata]
Clethrobius comes
Euceraphis betulae group,
caerulescens
Glyphina betulae,
pseudoschrankiana
Hamamelistes betulinus
Hormaphis betulae
Monaphis antennata
Stomaphis takahashii
Symydobius kabae
B. populifolia Calaphis betulaecolens, flavabg,
(Gray Birch) leonardi
Callipterinella callipterus
Euceraphis sp. near betulae,
lineata
Hamamelistes spinosus
B. pubescens Betulaphis quadrituberculata
(Downy Birch) Calaphis betulicola, flava
Callipterinella callipterus,
minutissima
Clethrobius comes
Euceraphis punctipennis
Glyphina betulae,
pseudoschrankiana
Hamamelistes betulinus
Monaphis antennata
Stomaphis quercus
Symydobius oblongus
 Betula 93

B. pumila Calaphis betulaefoliae,

(American Low Birch) manitobensis
Hamamelistes spinosus
B. sandbergii Betulaphis quadrituberculatabg
Calaphis flavabg
Callipterinella minutissimabg
Euceraphis betulaebg
B. schmidtii Clethrobius comes
B. tatewakiana Clethrobius comes
B. turkestanica Betulaphis hissarica
Calaphis flava
B. utilis Betacallis querciphaga
(Indian Paper Birch) Betulaphis longicornis
Clethrobius dryobius
[Eutrichosiphon sp.]
Neobetulaphis chaetosiphon,
pusilla
Betula sp. [Clethrobius vermai]
Eutrichosiphum (Paratrichosiphon)
betulae
[Mesocallis pteleae}

Key to birch-feeding aphids

1 Antennae of aptera very short, less than 0.2 x BL, 3- or 4-segmented.
Antennae of alata 4- or 5-segmented with annular (ring-like) rhinaria ... 2
- Antennae of aptera and alata 5- or 6-segmented. Secondary rhinaria
circular, oval or transversely elongate but never completely encircling
antenna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2 Adult aptera aleyrodiform, on upper surface of leaf, with wax glands all
around body producing a white wax fringe. Hind wing of alata with only
one oblique vein (sometimes branched) . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Adult apterae in gall or pseudogall on underside of leaf, not aleyrodiform
(but aleyrodiform or coccidiform adults or immatures may occur as
overwintering stages on twigs). Hind wing of alata with 2 separate oblique
veins ........................................................ 4
3 Aleyrodiform adult only about 0.6 mm long, dark brown to black in life
with a fringe of separate, rod-like wax filaments. Alata with 3 antennal
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hormaphis hamamelidis
- Aleyrodiform adult 1.0-1.2 mm long, yellowish or greenish-brown in life
with fringing wax filaments fused into plates. Alata with 5 antennal
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hormaphis betulae
94 Betula

4 Antenna of aptera 3-segmented, ANT III thin, at its base less than half
the width of ANT I, and tapering gradually from base towards apex
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes betulinus
- Antenna of aptera usually 4-segmented, sometimes 5-segmented, with
ANT III at its base at least half the width of ANT I . . . . . . . . . . . . . 5
5 Aptera (from gall) with pigmented siphuncular pores . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes gibberi
- Aptera (from gall) without siphuncular pores . . . . . . . . . . . . . . . . . . . . 6
6 Galls comprising reddish to crimson corrugations of leaf lamina between
veins. (Apterae in life dark brownish to purple with white wax) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes spinosus
- Galls comprising cockscomb-like bags protruding on upperside of leaf.
(Apterae reddish-yellow in life) . . . . . . . . . . . Hamamelistes cristafoliae
7 ANT PT/BASE less than or about equal to 1.0 . . . . . . . . . . . . . . . . . . 8
- ANT PT/BASE clearly more than 1.0, and usually more than 1.5 (the
single exception is one species with very small apterae, Callipterinella
minutissima, which has PT/BASE in range 1.1-1.8) . . . . . . . . . . . . . 37
8 Rostrum longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Rostrum shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9 HT II shorter than maximum diameter of SIPH cone . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis radicicola
- HT II longer than maximum diameter of SIPH cone . . . . . . . . . . . . 10
10 HT II more than 2 x longer than 2nd segments of fore and mid-tarsi
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis takahashii
- HT II less than 1.5 x longer than 2nd segments of fore and mid-tarsi
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis quercus
11 ANT PT peg-like, less than 2 x its basal width. Eyes of aptera reduced
to 3 facets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- PT of antenna finger-like, more than 2 x its basal width. If apterae are
present then their eyes are multifaceted . . . . . . . . . . . . . . . . . . . . . . . . 15
12 Dorsum pale. Body hairs inconspicuous. SIPH absent . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus ephemeratus
- Dorsum pigmented, hairs conspicuous. SIPH present as pores on small
cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Dorsal and lateral body hairs of aptera long and fine . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphina setosa
- Dorsal and lateral body hairs of aptera thick and spine-like . . . . . . 14
 Betula 95

14 Dorsum adorned with short wrinkles, sometimes extending into a

reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphina pseudoschrankiana
- Dorsum with wart-like cuticular ornamentation .... Glyphina betulae
15 Anal plate with a medial cleft, dividing into two lobes . . . . . . . . . . . 16
- Anal plate entire, rounded or with a slight medial indentation, but not
clearly bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
16 CAUDA of aptera with a very clear constriction, separating off a distinct
terminal knob . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- CAUDA of aptera semicircular or triangular, without a clear constriction
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
17 Antennae longer than body. SIPH with a single long hair . . . . . . . . 18
- Antennae much shorter than body. SIPH with or without hairs . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18 Knob of CAUDA elongate, about 2 x as long as wide. ANT VI (including
PT) less than 0.5 x ANT III . . . . . . . . . . . . . . . . . . . Taioa chuansiensis
- Knob of CAUDA only a little longer than wide. ANT VI almost as long
as ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taioa indica
19 SIPH with one or more capitate hairs arising from it posteriorly. ABD
TERG 1-3 of aptera with minute spinopleural hairs (8-10µm long)
. . . . . . . . . . . . . . . . . . . . . . . . . . Neobetulaphis chaetosiphon/immaculata
- SIPH without any hairs, and of alata with or without a hair. ABD TERG
1-3 of aptera with thick spinopleural hairs, expanded apically, 20-60µm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Aptera with ANT VI BASE more than 0.5 x ANT III. Longest
spinopleural hairs on ABD TERG 1-3 are 40-60 µm long . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neobetulaphis alba
- Aptera with ANT VI BASE less than 0.5 x ANT III. Longest spino-
pleural hairs on ABD TERG 1-3 are 20-30µm l o n g . . . . . . . . . . . . . . . .
........................................... Neobetulaphis pusilla
21 Aptera with all dorsal body hairs long and capitate (Fig. 31 A) .... 22
- Aptera with spinal and pleural hairs, or at least those on ABD TERG 1-4,
inconspicuous, much shorter than posterior marginal hairs and those on
ABD TERG 8 (Figs 31B-D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
22 SIPH of aptera densely spinulose . . . . . . . . . . . . . . Betulaphis hissarica
- SIPH of aptera smooth or weakly imbricated . . . . . . . . . . . . . . . . . . . 23
23 R IV+V shorter than HT II . . . . . . . . . . . Betulaphis quadrituberculata
- R IV+V longer than HT II . . . . . . . . . . . . . . . . . . Betulaphis longicornis
96 Betula

Fig. 31. Hairs on ABD TERG 2 and 3 of apterae of A, Betulaphis quadrituberculata (longer-haired form,
B, B. brevipilosa, C, B. pelei, D, B. quadrituberculata (shorter-haired form).

24 Aptera with spinal and pleural hairs on ABD TERG 7 as short and
inconspicuous as those on more anterior tergites. Marginal hairs on thorax
and ABD TERG 1-4 also very short. Dorsal abdominal hairs only long
and capitate on ABD TERG 8, and lateral hairs only long and capitate
on ABD TERG 5-8. Dorsal cuticle very wrinkled (Fig. 3IB) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betulaphis brevipilosa
- Aptera with at least some of spinal and pleural hairs on ABD TERG 5-7
long and capitate, in addition to marginal hairs and those on ABD TERG
8. Dorsal cuticle smooth or somewhat wrinkled . . . . . . . . . . . . . . . . . 25
25 R IV+V only about 0.06mm long, about as long as its basal width, less
than 0.6 x ANT VI BASE . . . . . . . . . . . . . . . . . . . . Betulaphis japonica
- R IV+V more than 0.07 mm long, more than 0.7 x ANT VI BASE
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Aptera with marginal hairs on ABD TERG 1-3 shorter than or about as
long as basal diameter of ANT III (Fig. 31C) . . . . . . . Betulaphis pelei
- Aptera with marginal hairs on ABD TERG 1-3 mostly much longer than
basal diameter of ANT III (Fig. 31D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . Betulaphis quadrituberculata (short-haired form)
 Betula 97

Fig. 32. Antennae of A, Monaphis antennata, B, Symydobius intermedius, C, S. kabae, D, S. oblongus.

27 CAUDA broadly rounded, without a trace of a constriction. Viviparae

both apterous (or brachypterous) and alate, with 2-3 broad white bands
on antennae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- CAUDA with a constriction separating the terminal part as a knob.
Viviparae all full-winged, with antennae not contrastedly banded . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
28 ANT PT/BASE about 1.0. Alata with wing veins heavily black-bordered
......................................... Symydobius americanus
- ANT PT/BASE clearly less than 1.0. Wing veins of alata with or without
black borders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 ANT III with 12-22 circular or oval secondary rhinaria spaced out along
length of segment (Fig. 32B). Wing veins of alata black-bordered ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symydobius intermedius
- ANT III with 16-35 transverse oval to elongate oval secondary rhinaria
closely set on basal 0.5-0.75 of segment (Fig. 32C). Wing veins of alata
not black-bordered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 Abdominal marginal sclerites each bearing 1-7 circular, transparent
tubercles (Fig. 29C). Body, legs and antennae clothed in numerous long,
fine hairs, those on ANT III and tibia being mainly longer than the
diameters of their respective segments . . . . . . . . . . . Symydobius kabae
- Abdominal marginal sclerites without tubercles. Hairs shorter and not so
fine, those on ANT III (Fig. 32D) and tibia mostly not exceeding diameters
of their respective segments . . . . . . . . . . . . . . . . . . Symydobius oblongus
31 Hairs on ANT III long and fine, 2 or more x longer than basal width of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
- Length of hairs on ANT III at most equal to basal diameter of the segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
98 Betula

32 SIPH bearing several hairs near base . . . . . . . . . . Clethrobius dryobius

- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33 Tibiae wholly black . . . . . . . . . . . . . . . . . . . . . . . . . . . . Clethrobius comes
- Tibiae at least partly pale . . . . . . . . . . . . . . . . . . . . . . Clethrobius vermai
34 Thorax with pale brown to black dorsal markings, but without a trace of
pigment ventrally. ANT VI including PT similar in length to or only a
little longer than ANT V . . . . . . . . . . . . . . . . . . . . . . . Euceraphis lineata
- Thorax similarly pigmented dorsally and ventrally. ANT VI including PT
rarely up to 0.75 x ANT V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35 Tibiae not uniformly dark; if mainly dark then the palest part is in the
middle and the distal part is scabrous and completely black . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euceraphis betulae group
- Tibiae wholly and rather uniformly dark . . . . . . . . . . . . . . . . . . . . . . . 36
36 Sensoriated part of ANT III extending up to 0.5 of length of segment,
leaving the distal, unsensoriated part clearly shorter than ANT IV ...
.............................................. Euceraphis mucida
- Sensoriated part of ANT III extending about 0.33 of length of segment,
leaving the distal, unsensoriated part about equal in length to ANT IV
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euceraphis ontakensis
37 ANT PT/BASE 7.0-9.0, with a very small primary sensorium between PT
and base of ANT VI. Secondary rhinaria circular and scattered on ANT
III, not in a row (Fig. 32A) . . . . . . . . . . . . . . . . . . . Monaphis antennata
- ANT PT/BASE not more than 3.0, with primary sensorium of normal
size. Secondary rhinaria on ANT III of alata oval or elongate oval and
more-or-less in a row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 SIPH long and tubular, with numerous long hairs . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum betulae
- SIPH in form of small truncate cones, without hairs . . . . . . . . . . . . 39
39 Alata with ANT PT/BASE less than 1.5. Apterous vivipara very small,
BL less than 1.5mm, with antennae 5-segmented, less than 0.5 x BL
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callipterinella minutissima
- Alata with ANT PT/BASE more than 1.5. Apterous vivipara, if present,
with BL greater than 1.5 mm and 6-segmented antennae at least 0.6 x BL
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40 Head and thorax deeply pigmented, and a pattern of dorsal abdominal
markings on more than two tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
- Head and thorax pale or only lightly pigmented, or pale with distinctive
dark markings, and dorsal abdomen with any dark markings restricted to
two tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
 Betula 99

41 Abdomen with separate dark dorsal transverse bars on all tergites ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callipterinella calliptera
- Abdomen with a large dark quadrate patch on ABD TERG 4 to 6
....................................... Callipterinella tuberculata
42 Head with a black transverse band across the venter between the eyes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
- Head without a ventral transverse band . . . . . . . . . . . . . . . . . . . . . . . . 46
43 Marginal tubercles on ABD TERG 2-5 or 4-5, and whole of SIPH,
pigmented. R IV+V less than 1.2 x HT II . . . . . . . . . . . . . . . . . . . . . . 44
- Marginal tubercles on ABD TERG 2-4 pale, those on 5 either pale or
pigmented. R IV+V at least 1.25 x HT II . . . . . . . . . . . . . . . . . . . . . . 45
44 Head and prothorax with a dark dorsal longitudinal median stripe.
Forewing veins evenly pigmented . . . . . . . . . . . . . Betacallis querciphaga
- Head and prothorax without a dorsal longitudinal median stripe. Media
of forewing pale, but Cu1b and distal part of Cu la thickly bordered with
fuscous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betacallis odaiensis
45 ANT III with 17-25 secondary rhinaria, on basal 0.5-0.7 of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Betacallis sikkimensis
- ANT III with 37-45 secondary rhinaria, extending more than 0.75 of
length of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . Betacallis luminiferus
46 Alata with a distinctive pattern of dark markings dorsally; longitudinal
stripes on head and thorax, and transverse bars or patches on abdomen
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
- Alata without a distinctive pattern of dark dorsal markings . . . . . . 48
47 Dorsal abdomen with separate double black bars on ABD TERG 4
and 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis betulella
- Dorsal abdomen with a large black quadrate patch on ABD TERG 4
and 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis neobetulella
48 Knob of CAUDA elongate, longer than wide . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis betulaefoliae
- Knob of CAUDA globular, no longer than its maximum width ... 49
49 Apterous morph . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
- Alate morph ................................................ 55
50 Most proximal secondary rhinarium on ANT III located near base of
segment, no further from base of segment than the length of ANT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
100 Betula

- Rhinaria on ANT III placed towards middle of segment, most proximal

one about as far from base as the combined lengths of ANT I+ II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
51 R IV+V 0.09-0.105 mm, less than 0.8 X HT II, and bearing only 2-4
accessory hairs besides the 3 subapical pairs ... Calaphis viridipallida
- R IV+V at least 0.11 mm long, more than 0.8 x HT II, with 4-8 accessory
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis flava
52 Antennae and legs wholly pale. Dorsal and lateral body hairs short and
inconspicuous, except on frons and posterior abdomen . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis manitobensis
- Antennae with dark apices to antennal segments, tibiae and tarsi. Dorsal
and lateral body hairs conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
53 Dorsal abdominal hairs arising from lightly pigmented tubercles. ANT III
only as long as, or a little longer than, head width across eyes . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis arctica
- Dorsal abdominal hairs often arising from tubercles, but these are
unpigmented. ANT III at least 1.5 x head width across eyes . . . . . 54
54 SIPH usually dark brown distally. R IV+V more than 0.125 mm long, as
long as or longer than HT II, with 14-23 accessory hairs. ANT III with
5-14 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis betulicola
- SIPH usually pale, occasionally a little darker at flange. R IV+V less than
0.12 mm long, shorter than HT II, with 4-7 accessory hairs. ANT III with
3-7 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis coloradoensis
55 SIPH with apical halves dark brown to black . . . . . . Calaphis betulicola
- SIPH pale, rarely a little dusky at flange . . . . . . . . . . . . . . . . . . . . . . . 56
56 ANT III with 4-8 secondary rhinaria . . . . . . . . Calaphis coloradoensis
- ANT III with 8-21 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . 57
57 Tibiae dark, forewing veins heavily bordered with fuscous . . . . . . . 58
- Tibiae pale except at base and apex, forewing veins dark but not heavily
bordered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
58 R IV+V usually more than 0.14 mm long, as long as or a little longer than
HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis betulaecolens
- R IV+V usually less than 0.14mm long, shorter than HT II . . . . . . 57
59 Hairs on ANT III very short, all less than 0.5 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis similis
- Hairs on ANT III mainly more than 0.5 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis leonardi
 Brachychiton 101

60 R IV+V 0.09-0.105 mm, less than 0.8 x HT II, and bearing only 2-4
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis viridipallida
- R IV+V at least 0.11 mm long, more than 0.8 x HT II, and bearing 4-8
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis flava

BISCHOFIA Euphorbiaceae

Bischofia javanica Toxoptera odinae

BIXA Bixaceae

Bixa orellana Neomyzus circumflexus

Sinomegoura citricola
(Use key to polyphagous aphids, p. 532)

BOMBAX Bombacaceae

Bombax buonopozense Toxoptera citricidus

Bombax ceiba Aphis gossypii
Toxoptera citricidus
(Use key to polyphagous aphids, p. 532)

BRACHYCHITON Sterculiaceae

Brachychiton spp. Myzus persicae

Sitobion sp.
102 Brachystegia

BRACHYSTEGIA Leguminosae

Brachystegia spp. Aphis craccivora

Aphis gossypii
(Use key to polyphagous aphids, p. 532)

BRIDELIA Euphorbiaceae

Host Plant List

Bridelia atroviridis Sitobion sp.
B. bridelifolia Sitobion halli
B. mollis [Aphis craccivora]
B. monoica Greenidea brideliae
Toxoptera citricidus
B. ovata Sinomegoura citricola
Toxoptera citricidus
B. tomentosa Greenidea brideliae, [sp. (Noordam,
in prep.)]
Greenideoida elongata,
{Greenideoida sp. (Noordam, in
prep.)]
Bridelia sp. Aphis gossypii
Schoutedenia ralumensis

Key to aphids on Bridelia

1 SIPH pores on very broad cones. ABD TERG 7 with a pair of long,
pointed, backward projections . . . . . . . . . . . . . Schoutedenia ralumensis
- SIPH tubular. ABD TERG 7 without long projections . . . . . . . . . . . . 2
2 SIPH with numerous long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Antenna 6-segmented, with ANT PT/BASE more than 2.0. SIPH with
a reticulate pattern of transverse striae over their entire length. CAUDA
with a terminal papilla . . . . . . . . . . . . . . . . . . . . . . . . Greenidea brideliae
 Buchania 103

Antenna 5-segmented, with ANT PT/BASE about 1.0. SIPH without a

reticulate pattern. CAUDA semicircular, without a papilla . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenideoida elongata
4 Antennal tubercles weakly developed. CAUDA tongue-shaped, rounded
at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Antennal tubercles moderately to well developed. CAUDA finger-like,
tapering distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 Stridulatory apparatus present (Fig. 122). CAUDA as black as SIPH, with
over 20 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera citricidus
No Stridulatory apparatus. CAUDA paler than SIPH, with 4-7 hairs
. . . . . . . . . . . . . . . . . . Aphis gossypii (or try key to polyphagous aphids)
6 SIPH about 0.2 x BL, similar in length to CAUDA, without any poly-
gonal reticulation (Fig. 121B) . . . . . . . . . . . . . . . . Sinomegoura citricola
- SIPH 0.25-0.35 x BL, clearly longer than CAUDA, with a subapical zone
of polygonal reticulation . . . . . . . . . . . . . . . Sitobion halli, Sitobion sp.

BROSIMUM Moraceae

Brosimum gaudichaudii Toxoptera aurantii

BROUSSONETIA Moraceae

Broussonetia papyrifera Aphis gossypii

BUCHANIA Anacardiaceae

Buchanania latifotia Formosaphis micheliae

104 Buddleja

BUDDLEJA Buddlejaceae

Buddleja spp. Aphis gossypii, spiraecola,

verbasci
Macrosiphum euphorbiae
[Mollitrichosiphon
(Metatrichosiphon) buddlejae}
Myzus antirrhinii, ornatus

Key to aphids on Buddleja

- Antennal tubercles poorly developed. SIPH black, tapering, curved
outward distally. R IV+V elongate, much more than 2 x its width at base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis verbasci
- Antennal tubercles poorly or well developed. SIPH pale, dusky or black.
R IV+V of normal length, about 2 x its width at base . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

BURSERA Burseraceae

Bursera simaruba Aphis gossypii

BUTEA Leguminosae

Butea spp. Aphis craccivora

Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

BUXUS Buxaceae

Buxus harlandi Myzus persicae

B. sempervirens Myzus persicae
 Calocedrus 105

CAESALPINIA Leguminosae

Aphis spiraecola
Myzus persicae
(Use key to polyphagous aphids, p. 532)

CALAMUS Palmae

Cerataphis ?variabilis
(For palm aphids see B & E, 1984)

CALLIGONUM Polygonaceae

Brachyunguis calligoni, harmalae,

plotnikovi
SIPH very small, about 0.03 mm long and 0.03 mm broad at base, and
only about 0.2-0.25 x CAUDA . . . . . . . . . . . . . Brachyunguis calligoni
- SIPH 0.06-0.08 mm long and 0.06-0.08 mm broad at base, about
0.5-0.6 x CAUDA . . . . . . . . . . . . . . Brachyunguis harmalae/plotnikovi

CALLITRIS Cupressaceae

Callitris spp. Cinara (Cupressobium) cupressi,

tujafilina
Illinoia morrisoni
(Use key to aphids on Cupressus, p. 161)

CALOCEDRUS Cupressaceae

Calocedrus decurrens Illinoia morrisoni

106 Calodendrum

CALODENDRUM Rutaceae

Calodendrum capense Toxoptera aurantii, citricidus

(Cape Chestnut)
(Use key to polyphagous aphids, p. 532)

CALOPHYLLUM Guttiferae

Calophyllum antillanum Toxoptera aurantii

C. inophyllum Aphis gossypii
(Use key to polyphagous aphids, p. 532)

CALOTROPIS Asclepiadaceae

Calotropis gigantea Aphis nerii

C. procera Aphis craccivora, nerii
Brachyunguis calotropicus,
harmalae, ushari
Myzus persicae

Key to aphids on Calotropis

I Antennal tubercles well developed, convergent in dorsal view (Fig. 42I).
SIPH tubular, slightly swollen on distal half . . . . . . . . Myzus persicae
Antennal tubercles weakly developed. SIPH tapering or conical .... 2
2 ANT PT/BASE a little less than 1. SIPH pale, truncated cones, much
shorter than CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- ANT PT/BASE much more than 1. SIPH dark, longer than CAUDA
............................................................. 4
3 SIPH 0.04-0.06 mm long, 0.33-0.40 x CAUDA . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyunguis calotropicus/ushari
- SIPH 0.06-0.08 mm long, about 0.5-0.6 x CAUDA . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyunguis harmalae
 Carissa 107

4 Dorsum of aptera usually with an extensive black patch. CAUDA pointed,

with 4-9 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
Dorsum of aptera without any dark markings. CAUDA blunt, with 9-19
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis nerii

CAMELLIA Theaceae

Camellia spp. Greenidea (Trichosiphum) sp.

(B. Agarwala, pers. comm.)
Sinomegoura citricola
Toxoptera aurantii, citricidus
[Aphis camellicola del Guercio
(nomen dubium)]
(Use key to polyphagous aphids, p. 532)

CANARIUM Burseraceae

Canarium album Sinomegoura citricola

Toxoptera odinae
(Use key to polyphagous aphids, p. 532)

CARISSA Apocynaceae

Carissa edulis Sitobion sp.

Toxoptera aurantii
C. grandiflora Aphis spiraecola
Toxoptera aurantii
C. macrocarpa Myzus persicae
(Use key to polyphagous aphids, p. 532)
108 Carpinus

CARPINUS Hornbeams Corylaceae

Host Plant List

Carpinus betulus Myzocallis carpini
C. caroliniana Macrosiphum (Neocorylobiuni)
carpinicolens
C. caucasica Myzocallis carpini
C. cordata Mesocallis sawashibae
Myzocallis coryli
[Tinocallis zelkowae]
C. coreana Mesocallis sawashibae
C. erosa Mesocallis sawashibae
C. faginea Greenidea (Trichosiphurri)
carpinicola
[Mesocallis alnicola]
C. japonica Mesocallis sawashibae
Neochromaphis carpinicola
Pterocallis montana
C. laxiflora Macrosiphum (Neocorylobium)
corylicola
Neochromaphis carpinicola,
?coryli
C. tschonoskii Neochromaphis carpinicola
Stomaphis carpini
C. viminea Greenidea (Trichosiphum)
carpinicola
Carpinus spp. Greenidea (Trichosiphum) carpini
Kaltenbachiella carpinicola
Mesocallis pteleae

Key to aphids on Carpinus

1 SIPH long and tubular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- SIPH short, conical or ring-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 SIPH with numerous long fine hairs. CAUDA broad, rounded .... 3
- SIPH without hairs, but with an apical zone of polygonal reticulation.
CAUDA long and finger-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
 Carpinus 109

3 R IV+V c. 1.8 x HT II . . . . . . . . . . . Greenidea (Trichosiphum) carpini

- R IV+V 1.4-1.6 x H II . . . . . . . Greenidea (Trichosiphum) carpinicola
4 ANT III mainly smooth, with longest hairs more than 0.5 x basal
diameter of segment. R IV+V 1.3-1.9 x HT II . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) corylicola
- ANT III imbricated throughout, with longest hairs less than 0.5 x basal
diameter of segment. R IV+V 1.1-1.3 x HT II . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) carpinicolens
5 Rostrum longer than body. SIPH placed on large, pigmented hairy cones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis carpini
- Rostrum much shorter than body. SIPH small, conical, truncate or
ring-like ..................................................... 6
6 CAUDA broadly rounded, anal plate entire or only slightly indented
(apterae and immatures in leaf galls) . . . . . Kaltenbachiella carpinicola
- CAUDA with a knobbed apex, anal plate deeply bilobed. Adult viviparae
all alate, with one exception (Mesocallis sawashibae) . . . . . . . . . . . . . . 7
7 Wings of alata with patches of pigmentation . . . . . . . . . . . . . . . . . . . . . 8
- Wings of alata without pigmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Longest hairs on ANT III less than 3 x diameter of segment at base.
R IV+V 1.0-1.3 x HT II . . . . . . . . . . . . . . . Neochromaphis carpinicola
- Longest hairs on ANT III more than 3 x basal diameter of segment.
R IV+V 1.4-1.6 x HT II . . . . . . . . . . . . . . . . . . . . Neochromaphis coryli
9 ANT PT/BASE 0.2-0.45 . . . . . . . . . . . . . . . . . . . . . . Pterocallis montana
- ANT PT/BASE 0.5 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 ANT PT/BASE at least 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- ANT PT/BASE 1.2 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 R IV+V 1.4-1.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis coryli
- R IV+V 0.9-1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . Myzocallis carpini
12 Front of head and ANT III dark. R IV+V 1.2-1.4 X HT II. All viviparae
alate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesocallis pteleae
- Front of head and ANT III pale. R IV+V 0.8-1.0 x HT II. Viviparae
apterous or alate . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesocallis sawashibae
110 Garya

CARYA Hickories Juglandaceae

The aphids of pecan (Carya illinoensis) were keyed in Blackman and Eastop
(1984). The present key is extended to cover other Carya species, but there
are certain difficulties. The hickory phylloxerids (Phylloxera spp.), with more
than 30 available names, are impossible to key in the present state of know-
ledge. Pergande (1904) provided a classification and keys based on the galls.
Many species of the three North American genera Monellia, Monelliopsis and
Protopterocallis are morphologically similar, yet most are recorded from
several Carya species, so a single key is necessary with some rather difficult
and possibly unreliable discriminants within these genera.

Host Plant List

Carya amara = C. cordiformis
C. aquatica Melanocallis caryaefoliae
(Water Hickory) Monellia caryella
C. cathayensis Kurisakia sinocaryae
(Chinese Hickory, Chinese Walnut)
C. cordiformis Melanocallis caryaefoliae
(Bitternut Hickory) Monellia caryella, hispida
[Phylloxera caryaemagna, conica,
foveata, globosa, minima]
Protopterocallis fumipenella
C. glabra (= C. porcina) Longistigma caryae
(Pignut Hickory) Melanocallis caryaefoliae
Monellia caryella, hispida,
microsetosa
[Phylloxera caryaecaulis,
caryaefoliae, caryaeren,
caryaesemen, caryaesepta var.
perforans, pilosula]
Protopterocallis fumipenella
C. glabra var. odorata = C. ovalis
C. illinoensis [Chromaphis californica]
(Pecan) Longistigma caryae
Melanocallis caryaefoliae
Monellia caryella, medina
Monelliopsis nigropunctata,
pecanis
Phylloxera devastatrix, notabilis,
perniciosa(?), russellae, texana
 Carya 1ll

C. laciniosa (= C. sulcata) Melanocallis caryaefoliae

(Shellbark Hickory) Monellia caryella, microsetosa
C. ovalis Melanocallis caryaefoliae
(Red Hickory) Monellia hispida, microsetosa
C. ovata Melanocallis caryaefoliae
(Shagbark Hickory) Monellia caryella, costalis,
hispida, medina, microsetosa
Monelliopsis nigropunctata
[Phylloxera caryaefallax,
caryaeglobuli, caryaemagna,
caryaesepta, caryaevenae,
conica, depressa, subelliptica]
C. pallida Melanocallis caryaefoliae
(Sand Hickory) Protopterocallis pergandei
C. tomentosa Melanocallis caryaefoliae
(Mockernut Hickory) Monellia hispida, microsetosa
Monelliopsis nigropunctata
[Phylloxera caryaeavellana,
caryaescissa, caryaevenae,
castaneae, deplanata,
intermedia, perniciosa, picta,
rimosalis, symmetrica]
Protopterocallis, fumipenella,
gigantea, quadrata
Carya sp. [Phylloxera depressa, georgiana,
spinosa, spinuloida]
[Prociphilus caryae]

Key to aphids colonizing Carya spp

1 Antennae of adults 3- or 4-segmented . . . . . . . . . . . . . . Phylloxera spp.
- Antennae 5- or 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 BL more than 4 mm. Alata with pterostigma extending around tip of
forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- BL less than 3.5 mm. Alata with pterostigma stopping short of forewing
tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 CAUDA rounded, anal plate entire. SIPH cones bearing numerous long
hairs. ANT PT/BASE less than 0.4 . . . . . . . . . . . Kurisakia sinocaryae
- CAUDA knobbed, anal plate deeply bilobed. SIPH cones with at most
1-2 short hairs. ANT PT/BASE greater than 0.4 (all viviparae alate)
............................................................. 4
4 SIPH black, truncate, similar in length to ANT I (Fig. 33A). Head and
112 Carya
 Carya 113

thorax, and mid- and hind femora, all very dark, as are the well-developed
spinal and marginal abdominal tubercles ..... Melanocallis caryaefoliae
- SIPH pale and often inconspicuous, small, tuberculate. Head thorax and
all femora mainly pale or dusky, abdominal tubercles variably developed
and/or pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Forewing vein Cu1b ('anal' vein) darker than vein Cu1a, lightly bordered
with fuscous along most of its length. Rhinaria-bearing part of ANT III
only slightly thickened, rarely up to 1.5 x diameter of basal articulation
of segment (Fig. 33B). Embryos (seen through adult cuticle) with spinal
hairs on ABD TERG 5 displaced laterally, and spinal hairs on ABD TERG
6 displaced medially (Figs 34A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Forewing vein Cu lb as pale as Cu1a (except sometimes at ends). Rhinaria-
bearing part of ANT III conspicuously swollen, usually more than
1.5 x diameter of basal articulation of segment (Fig. 33C). Embryos with
spinal hairs on ABD TERG 1-6 in two longitudinal rows without any
lateral or medial displacement (Figs 34C-E) . . . . . . . . . . . . . . . . . . . . . 14
6 ANT PT/BASE 0.6-1.1. Small, delicate aphids, BL rarely more than
2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- ANT PT/BASE 0.3-0.6. More robust aphids, BL usually more than 2 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
7 ABD TERG 1-6 each with only one pair of spinal hairs, sometimes arising
from tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- ABD TERG 1-6 each with at least 4 spinopleural hairs, either as separate
spinal and pleural pairs, or with spinal hairs duplicated on flat tubercles
(bosses) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Pronotum, metanotum and ABD TERG 1-3 each with a pair of well-
developed tubercles, much longer than the flattened, pigmented bosses on
ABD TERG 4-6 (Fig. 33D). Embryos (seen through adult cuticle) with
spinal hairs on ABD TERG 2-4 long enough to overlap when directed
towards one another (Fig. 34A) . . . . . . . . . . . . Monelliopsis tuberculata
- Pronotum and metanotum without tubercles, and ABD TERG 1-3 with
flattened hair-bearing bosses like those on ABD TERG 4-6 (Fig. 33E).
Embryos with spinal hairs on ABD TERG 2-4 too short to overlap when
directed towards one another (Fig. 34B) . . . . . . . . . Monelliopsis caryae

Fig. 33. A, SIPH of Melanocallis caryaefoliae; B, ANT III of Monelliopsis caryae; C, ANT III of Monellia
caryella; D, tubercular bases of spinal hairs on ABD TERG 1-6 of Monelliopsis tuberculata; E, the same
for M. caryae; F, position of dark spot on hind femur of Monelliopsis bisetosa; G, the same for
M. nigropunctata; H, tubercular bases of spinal hairs on ABD TERG 2-4 of Monelliopsis nigropunctata;
I, the same for M. pecanis; J, hair-bearing tubercles on prothorax of Protopterocallis quadrata; K, the
same for P. pergandei; L, the same for P. gigantea; M, hair-bearing tubercles on ABD TERG 1 and 2 of
P. gigantea.
114 Carya
 Carya 115

9 ABD TERG 1-6 without pleural hairs, but with 2 spinal hairs on each
spinal boss. Hind femur with a black spot dorsally near apex (Fig. 33F)
............................................ Monelliopsis bisetosa
- ABD TERG 1-6 with separate spinal and pleural hairs on separate bosses.
Hind femur with a black spot on ventral side near apex (Fig. 33G) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Spinal tubercles (bosses) on ABD TERG 2-4 clearly larger than pleural
bosses (Fig. 33H) . . . . . . . . . . . . . . . . . . . . . . Monelliopsis nigropunctata
- Spinal bosses on ABD TERG 2-4 of similar size to pleural bosses
(Fig. 33I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monelliopsis pecanis
11 Pronotum with 2 posterior tubercles (Fig. 33J), and ABD TERG 1-4 each
with 4 tubercles . . . . . . . . . . . . . . . . . . . . . . . . . Protopterocallis quadrata
- Pronotum usually with at least 4 posterior tubercles (Figs 33K, L), and
ABD TERG 1-4 each usually with more than 4 tubercles . . . . . . . . 12
12 Pronotum with 6-7 posterior tubercles, in groups of 2-4 (Fig. 33K)
....................................... Protopterocallis pergandei
- Pronotum with 4-5 posterior tubercles (Fig. 33L) . . . . . . . . . . . . . . . . 13
13 Some of spinal hairs on ABD TERG 1-3 neotenic; as long as or longer
than the tubercles from which they arise, 45-80 µm long, expanded at tips
and very stout-based (Fig. 33M) . . . . . . . . . . . Protopterocallis gigantea
- Spinal hairs on ABD TERG 1-3 all shorter than the tubercles from which
they arise, less than 40 µm long, similar in size to hairs on pronotum
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protopterocallis fumipenella
14 ABD TERG 1-6 each with 3-8 fine, often very short hairs, with small
round pigmented spots at their bases, which are never tuberculate ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia microsetosa
- ABD TERG 1-6 each with only 2 spinal hairs, often on low tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Spinal hairs on ABD TERG 1-4 all short and fine, less than 40 µm
long, less than 1.5 x diameter of most swollen part of ANT III. ABD
TERG 1-6 of embryos (seen through adult cuticle) with very short spinal
hairs and long overlapping marginal hairs (Fig. 34C) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia caryella
- At least some of spinal hairs on ABD TERG 1-4 long, 40-80 µm,
1.5-3.2 x diameter of most swollen part of ANT III. If ABD TERG 1-6
of embryos have very short spinal hairs (Monellia hispida) then the
marginal hairs do not overlap . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Embryos (and 1st instars) with marginal hairs on abdominal segments
1-6 short, not overlapping between segments, and spinal hairs minute
(Fig. 34D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia hispida
116 Caryocar

- Embryos (and 1st instars) with marginal and spinal hairs all long, the
marginal hairs all overlapping between segments (Fig. 34E) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monellia medina

CARYOCAR Caryocaraceae

Caryocar brasiliense Aphis gossypii

CASEARIA Flacourtiaceae

Casearia arborea Toxoptera aurantii

C. decandra Aphis spiraecola
C. sylvestris Toxoptera aurantii
(See key to polyphagous aphids, p. 532)

CASSIA Leguminosae

Cassia corymbosa Aphis craccivora, fabae group

C. didymobotrya Aphis gossypii
C. siamea [Kurisakia yunnanensis]
Cassia sp. (African) Sitobion burundiense

CASTANEA Chestnuts Fagaceae

Host Plant List

Castanea ashei Longistigma caryae
Patchia virginiana
C. bungeana Castanocallis castanocallis, cereus,
margituberculatus
Myzocallis kuricola
 Castanea 117

C. crenata Aulacorthum magnoliae

(Japanese Chestnut) Cervaphis quercus
[Eutrichosiphum sp. (Noordam, in
prep.)]
Greenidea kuwanai, [spp.
(Noordam, in prep.)]
[Kurisakia onigurumii]
Lachnus tropicalis
Mollitrichosiphon tenuicorpus
Moritziella castaneivora
Myzocallis castanicola, kuricola
Tuberculatus (Orientuberculoides)
capitatus ssp. intermedius
C. dentata Myzocallis castaneae, castaneoides,
(American Chestnut) castanicola
Patchia virginiana
Phylloxera castaneae
C. formosana Eutrichosiphum tattakanum
C. henryi Myzocallis kuricola
C. mollissima Myzocallis kuricola
Phylloxera castaneae
C. nana Myzocallis castaneoides, nana,
tissoti
C, pubinervis Cervaphis quercus
Myzocallis kuricola
C. pumila Myzocallis castanicola
Patchia virginiana
Phylloxera castaneae
Tuberculatus annulatus
C. sativa Lachnus roboris, tropicalis
(Sweet Chestnut) Myzocallis castaneoides,
castanicola, castanicola ssp.
leclanti, kuricola
Phylloxera castaneae, [spinifera]
Thelaxes suberi
Tuberculatus annulatus

Key to aphids feeding on Castanea

1 Antennae 3-segmented (small insect, body pyriform, broadest anteriorly
............................................................. 2
- Antennae 5- or 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
118 Castanea

2 Many dorsal tubercles present on head, thorax and ABD TERG 1-6,
decreasing in size and number posteriorly . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moritziella castaneivora
- Only 4 dorsal tubercles present, confined to head . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera castaneae
3 SIPH tubular, several times longer than w i d e . . . . . . . . . . . . . . . . . . . . . 4
- SIPH truncate or broad, hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4 Body of aptera with very long, branched, hair-bearing lateral processes
(Fig. 92A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cervaphis quercus
- Body of aptera without hair-bearing processes . . . . . . . . . . . . . . . . . . . 5
5 SIPH without hairs. Antennal tubercles well developed, approx. parallel-
sided in dorsal view (Fig. 42J) . . . . . . . . . . . . . Aulacorthum magnoliae
- SIPH with numerous long fine hairs. Antennal tubercles undeveloped
............................................................. 6
6 Abdomen of aptera broadly rounded, uniformly brown pigmented. SIPH
of aptera with a heavily sclerotized, reticulate zone at the bases, and
CAUDA with a distinct median conical papilla (as in Fig. 92B) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) kuwanai
- Abdomen of aptera oval, less pigmented, at least with a pale anterio-
median area. SIPH of aptera without basal reticulation, and CAUDA
without a median papilla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 SIPH of aptera more than 0.7 x BL. Hind tibiae with transverse
stridulatory ridges (as in Fig. 27A) .... Mollitrichosiphon tenuicorpus
- SIPH of aptera less than 0.7 x BL. Hind tibiae without stridulatory ridges
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum tattakanum
8 ANT PT/BASE about 1.0 or more. Anal plate divided by a median cleft
into 2 lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- ANT PT/BASE 0.5 or less. Anal plate not bilobed . . . . . . . . . . . . . . 21
9 CAUDA triangular, apically pointed, without a constriction (Fig. 35A).
Body and appendages almost entirely brown-black . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patchia virginiana
- CAUDA with a constriction and a terminal knob. Body and appendages
mainly pale, sometimes with dark markings . . . . . . . . . . . . . . . . . . . . . 10
10 Head and pronotum with a longitudinal dark spinal stripe, and ABD
TERG 1-7 each with paired dark dorsal spinal as well as marginal sclerites
(Fig. 35E). (Wings not maculate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Head and pronotum without a spinal longitudinal stripe. If ABD TERG
1-7 have a complete set of paired dark dorsal sclerites then wings are
strongly maculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
 Castanea 119

Fig. 35. A, CAUDA oi Patchia virginiana; B, front of head and ANT I-II of Castaneomyzocallis
castaneae; C, the same for C. nanae; D, the same for C. castaneoides; E, dorsal body markings of
Myzocallis castanicola; F, marginal tubercles of ABD TERG 3-6 and SIPH of Castanocallis cereus;
6, spinal tubercles on ABD TERG 2 of Castanocallis castanocallis; H, the same for Myzocallis kuricola;
I, the same for Tuberculatus capitatus.

11 R IV with 4-10 accessory hairs. First tarsal segments with 5-6 ventral
hairs. Embryos with long hairs (55-70 µm) on ABD TERG 2 . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castanicola castanicola
- R IV with 11-18 accessory hairs. First tarsal segments with 6-8 (usually
7) ventral hairs. Embryos with hairs on ABD TERG 2 shorter (27-41 µm)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castanicola leclanti
12 Tibiae mainly brown to black, or at least brown on basal half. ANT
PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Tibiae mainly pale, sometimes dark apically. ANT PT/BASE less than 2.0
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
120 Castanea

13 Anterior frontal hairs of head long and finely pointed, 2 or more X basal
diameter of ANT III (Fig. 35B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Anterior frontal hairs less than 2 x basal diameter of ANT III, and
sometimes with capitate apices (Figs 35C, D) . . . . . . . . . . . . . . . . . . . . 15
14 Tibiae and ANT III-VI uniformly black . . . . . . . Myzocallis castaneae
- Tibiae dark at base but with paler middle region. ANT IV, V and BASE
VI each pale at base and dark at apex . . . . . . . . . . . . Myzocallis tissoti
15 Anterior frontal hairs 7-15 µm long, shorter than basal diameter of ANT
III, and capitate (Fig. 35C). ANT III-VI pale basally and dark apically,
and tibiae usually with paler middle region . . . . . . . . Myzocallis nanae
- Anterior frontal hairs 30-38 µm, longer than basal diameter of ANT III,
pointed or slightly capitate (Fig. 35D). ANT III-VI and tibiae often
completely dark . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castaneoides
16 Forewings with pigment extending between veins, or with veins at least
heavily bordered with pigment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Forewings not pigmented between veins . . . . . . . . . . . . . . . . . . . . . . . . 20
17 ANT III with 4-6 hairs, maximally about 1.5 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castanocallis margituberculatus
- ANT III with 7 or more hairs, including anteriorly directed long ones more
than 2 x basal diameter of the segment . . . . . . . . . . . . . . . . . . . . . . . . 18
18 ANT III with more than 20 hairs. Marginal tubercles each with 4-6
transparent noduli (Fig. 35F) . . . . . . . . . . . . . . . . . . Castanocallis cereus
- ANT III with less than 15 hairs. Marginal tubercles without any
transparent noduli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 ABD TERG 2 and 3 each with a pair of quadrate spinal tubercles, as long
as or longer than their basal widths (Fig. 35G). ANT III longer than head
width across (and including) eyes . . . . . . . . . Castanocallis Castanocallis
- Spinal tubercles on ABD TERG 2 and 3 if developed then rounded, and
shorter than their basal widths (Fig. 35H). ANT III shorter than head
width across eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis kuricola
20 Spinal tubercles on ABD TERG 1-3 all similarly developed, finger-like,
longer than their basal widths (Fig. 35I). Anteriorly directed hairs on front
of head and ANT I-III long and capitate. SIPH pale . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus capitatus
- Spinal tubercles only ever well developed on ABD TERG 3, and then
conical, shorter than their basal widths. Hairs on front of head and ANT
I-III short or very short, and pointed. SIPH black except at bases
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus annulatus
21 BL less than 2 mm. Antennae 5-segmented. CAUDA knobbed . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelaxes suberi
 Castanopsis 121

- BL more than 2.5 mm. Antennae 6-segmented. CAUDA broadly rounded

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 ABD TERG 8 unmarked or with only irregular pigment spots. Alata with
pterostigma extending around tip of forewing .... Longistigma caryae
- ABD TERG 8 with a thin transverse band. Alata with pigmented
(maculate) forewings and normal pterostigma . . . . . . . . . . . . . . . . . . . 23
23 ANT/PT BASE 0.40-0.68, usually more than 0.5. Alata with forewings
pigmented to base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus tropicalis
- ANT/PT BASE 0.3-0.5. Alata with clear area at base of forewing
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus roboris

CASTANOPSIS Fagaceae

Host Plant List

Castanopsis acuminatissima Eutrichosiphum pasaniae,
[Eutrichosiphum sp. (Noordam, in
prep.)]
[Greenidea sp. (Noordam, in
prep.)]
Nipponaphis javanica
C. argentea [Eutrichosiphum sp. (Noordam, in
prep.)]
[Greenidea spp. (Noordam, in
prep.)]
Nipponaphis brevipilosa
C. carlesii Metanipponaphis ?echinata
C. cuspidata Eutrichosiphum pasaniae, shiicola,
sinense
Greenidea okajimai
Lachnus tropicalis (or shiicola?)
Lithoaphis shiiae
Metanipponaphis cuspidatae,
shiicola
Mollitrichosiphon tenuicorpus
Neonipponaphis shiiae
[Nipponaphis distyliicola]
Reticulaphis shiiae
C. formosana Eutrichosiphum tattakanum
Mollitrichosiphon tenuicorpus
122 Castanopsis

C. histrix Metanipponaphis echinata

C. indica Eutrichosiphum pseudopasaniae
C. javanica Eutrichosiphum sinense
[Eutrichosiphum spp. (Noordam,
in prep.)]
[Greenidea spp. (Noordam, in
prep.)]
Nipponaphis multisetosa
C. sclerophylla Eutrichosiphum sclerophyllum
Metanipponaphis sp.
C. subacuminata Mollitrichosiphon tenuicorpus
C. submacula Eutrichosiphum pasaniae
C. tribuloides Metanipponaphis assamensis
Castanopsis sp(p). [Greenidea kunmingensis]
Lachnus shiicola
Metanipponaphis rotunda
Mollitrichosiphon elongatum
Nipponaphis querciphaga

Key to aphids feeding on Castanopsis

1 Aptera with head, thorax and ABD TERG 1 fused as prosoma. Legs and
antennae much reduced, often concealed beneath body. SIPH small or
absent. Antennae of alata with narrow ring-like sensoria . . . . . . . . . . 2
- Aptera of normal aphid form with well-developed legs and antennae.
SIPH evident, conical or tubular. Antennae of alata with circular or oval
sensoria ..................................................... 9
2 Aptera without SIPH; tarsi and claws rudimentary; dorsum strongly
reticulated, without pustules (Fig. 36A) . . . . . . . . . . . Reticulaphis shiiae
- Aptera with SIPH usually present as small pores or cones; tarsi and claws
developed; dorsum not reticulated, either mainly smooth or with
numerous transparent areas or raised pustules . . . . . . . . . . . . . . . . . . . . 3
3 Dorsal cuticle of prosoma mainly smooth except for areolation, pustules
only present around thoracic spiracles. Prosomal hairs long and thick with
somewhat tuberculate bases. ABD TERG 8 with 4 hairs . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis quercicola
- Dorsal cuticle of prosoma ornamented with numerous raised pustules or
translucent rounded or curved areas (Fig. 36B-E). Prosomal hairs short
or if long then fine. ABD TERG 8 with 2-10 hairs . . . . . . . . . . . . . . 4
4 Abdominal plate (ABD TERG 2-7) distinctly separate from prosoma
(Fig. 36B). ABD TERG 8 with 4-6 hairs ..... Neonipponaphis shiiae
 Castanopsis 123

E
C
Fig. 36. A, margin of prosoma of aptera of Reticulaphis shiiae; B, dorsal view of aptera of
Neonipponaphis shiiae, with detail of cuticular ornamentation of prosoma; C, the same for
Metanipponaphis sp.; D, cuticular ornamentation of prosoma of aptera of Lithoaphis shiiae; E, the same
for Nipponaphis brevilosa.

- Abdominal plate fused with or not distinctly separate from prosoma,

at least in median area (Fig. 36C). ABD TERG 8 with 2-10 hairs ....
............................................................. 5
5 ABD TERG 8 with 2-4 hairs. Prosoma dorsally ornamented with densely
packed, evenly distributed, rather uniform-sized, circular or oval pustules
(Fig. 36C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metanipponaphis spp.
- ABD TERG 8 with 4-6 hairs. Prosoma with polygonal, angular or
irregular dorsal ornamentation; if pustulate, then the pustules are of
irregular size and shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Prosoma strongly convex dorsally, with irregular translucent branched or
curved ornamentation (Fig. 36D). ABD TERG 8 with 6 hairs . . . . . . . .
............................................... Lithoaphis shiiae
124 Castanopsis

- Prosoma somewhat flattened dorsally, with variable pustulate or poly-

gonal ornamentation (e.g. Fig. 36E). ABD TERG 8 with 4 hairs ..... 7
7 Prosomal hairs only 10-20 µm long, and those on ABD TERG 8 only
35-40 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis brevipilosa
- Prosomal hairs 40-125 µm long, and those on ABD TERG 8 65-160 µm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Prosoma with only 6-7 hairs (2-3 spinal, 4 marginal) on each thoracic
tergite, and 4 hairs (2 spinal, 2 marginal) on ABD TERG 1 . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis javanica
- Prosomal hairs much more numerous; 30-70 per tergite . . . . . . . . . . . .
......................................... Nipponaphis multisetosa
9 SIPH tubular, bearing numerous long hairs . . . . . . . . . . . . . . . . . . . . . 10
- SIPH truncate or broadly conical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
10 SIPH of aptera with heavily sclerotized reticulate zone at base, and
CAUDA with a terminal papilla. Head capsule of alata (including eyes)
clearly more than 2 x as wide as long dorsally . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) okajimai
- SIPH of aptera without reticulation, CAUDA rounded without a papilla.
Head of alata (including eyes) about 2 x as wide as long dorsally ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Hind tibiae with transverse (stridulatory?) ridges (e.g. Fig. 27A) . . . 12
- Hind tibiae without transverse ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12 Antennae 5- or 6-segmented. SIPH of aptera less than 0.5 x BL . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphon elongatum
- Antennae 6-segmented. SIPH of aptera more than 0.7 x BL . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphon tenuicorpus
13 Dorsum of aptera without spinules ...... Eutrichosiphum tattakanum
- Dorsum of aptera with numerous irregular transverse rows of minute
spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Aptera with R IV+V 1.5-1.7 x HT II, and SIPH longer than maximum
body width. SIPH of alata more than 1.5 x ANT III . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum shiicola
- Aptera with R IV+V 1.8-2.5 x HT II, and SIPH shorter than maximum
body width. SIPH of alata less than 1.5 x ANT III . . . . . . . . . . . . . 15
15 Dorsum of aptera pale, SIPH black distally, pale at base, and at least
0.7 x maximum body width . . . . . . . . . . . . . . . . Eutrichosiphum sinense
- Dorsum of aptera pale or dark brown, SIPH wholly brown-black and
0.5-0.6 x maximum body width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
 Catalpa 125

16 Dorsum of aptera brown-black, R IV+V 1.9-2.3 x HT II . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pasaniae
- Dorsum of aptera pale, sometimes dark at margins; R IV+V 2.2-2.5 x
HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pseudopasaniae
17 BL more than 3mm, ANT PT/BASE about 0.5. Forewings of alata
extensively dark-pigmented with two pale patches . . . . . . . . . . . . . .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus tropicalis
- BL less than 2.5 mm, ANT PT/BASE about 1.0. Forewings of alata often
with broad bands of pigment bordering veins, but with clear areas in
between . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis (Nippocallis) kuricola

CASTANOSPERMUM Leguminosae

Castanospermum australe Betacallis querciphaga

(Australian Chestnut) Mollitrichosiphon tenuicorpus

CASUARINA Casuarinaceae

Casuarina equisetifolia Aphis gossypii

[Lachnus ?tropiealis]
Myzus persicae
C. huegeliana Myzus persicae
C. torulosa Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

CATALPA Bignoniaceae

Catalpa spp. Aphis catalpae, fabae, gossypii,

nerii, spiraecola
Aulacorthum solani
Macrosiphum euphorbiae
Myzus ornatus, persicae
Toxoptera aurantii
126 Cecropia

Key to aphids on Catalpa

1 Antennal tubercles well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
Antennal tubercles weakly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH and CAUDA both very dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
- CAUDA pale or dusky, SIPH pale, dusky or dark . . . . . . . . . . . . . . . 3
3 Alata usually with no secondary rhinaria on ANT IV (rarely 1-2). SIPH
of aptera uniformly dusky to dark, 1.3-2.5 x CAUDA . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
Alata with 1-4 rhinaria on ANT IV. Aptera with SIPH usually paler
towards base, 1.10-1.35 x CAUDA . . . . . . . . . . . . . . . . . Aphis catalpae

CECROPIA Cecropiaceae

Cecropia peltata Aphis gossypii, spiraecola

(Use key to polyphagous aphids, p. 532)

CEDRELA Meliaceae

Cedrela odorata Toxoptera aurantii

(West Indian or Mexican Cedar)

CEDRUS Cedars . Pinaceae

Cedrus atlantica Cinara (Cedrobium) laportei

Cinara cedri
C. deodora Cinara (Cedrobiuni) laportei
Cinara confinis, curvipes, indica
Illinoia morrisoni
C. libani Cinara (Cedrobiuni) laportei
 Cedrus 127

Fig. 37.A, dorsal hair of aptera of Cinara (Cedrobium) laportei; B, HT I of Cinara cedri.

Key to aphids on Cedrus

1 ANT PT/BASE more than 1. SIPH long and tubular, swollen distally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni
- ANT PT/BASE less than 1. SIPH are broad hairy cones . . . . . . . . . 2
2 Antenna 5-segmented. Dorsal hairs of aptera club-shaped, ornamented
with numerous barbules (Fig. 37A) . . . . . Cinara (Cedrobium) laportei
- Antenna 6-segmented. Dorsal hairs normal, pointed . . . . . . . . . . . . . . 3
3 Hairs on body and appendages short; those on ANT III maximally about
as long as basal diameter of segment . . . . . . . . . . . . . . . Cinara curvipes
Hairs mostly long; longest hairs on ANT III maximally more than
2 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 BL 3.0mm or less. Dorsal length of HT I more than 1.5 x basal width
(Fig.37B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara cedri
BL more than 3.0mm. Dorsal length of HT I shorter than basal width
............................................................. 5
5 Aptera with HT II less than 4 x HT I. ANT III with more than 40 long
hairs, very few of these less than 2 x basal diameter of segment. BL
3.8-7.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara confinis
- Aptera with HT II 4 or more x HT I. ANT III with les.s than 30 hairs
of very variable length, often less than 2 x basal diameter of segment. BL
about 3.3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara indica
128 Celtis

CELTIS Ulmaceae

Celtis africana Toxoptera aurantii

C. aurantiaca [Aphis gossypii]
Shivaphis szelegiewiczi
C. australis Shivaphis catalpinari, celti,
celticola
Toxoptera aurantii
C. caucasica = C. australis
C. cinnamomia Shivaphis celti
C. eriocarpa = C. australis
C. formosana Aulacorthum solani
C. jessoensis Prociphilus oriens
Shivaphis celti
C. julianae Shivaphis celti, hanzhouensis
C. nervosa Shivaphis celti
C. sinensis [Aphis spiraecola]
(Enoki) Aulacorthum magnoliae
Paracolopha morrisoni
Shivaphis celti
Stomaphis yanonis
Toxoptera aurantii, celtis
C. tetrandra Shivaphis celti
Sumatraphis celti
C. triandra Shivaphis celti

Key to aphids on Celtis

I ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2 (Alata emerging in spring from galls or leaf-nests.) Media of forewing
unbranched .................................................. 3
- (Apterae and alatae free-living.) Media of forewing of alata once- or
twice-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 (Alata from closed gall.) BL less than 3 mm. Hind wing with one oblique
vein. Secondary rhinaria on ANT III-VI, many of them almost completely
encircling antenna . . . . . . . . . . . . . . . . . . . . . . . . . Paracolopha morrisoni
 Celtis 129

Fig. 38. A, posterior part of abdomen (right side) of Sumatraphis celti; B, CAUDA and anal plate of
Shivaphis catalpinari; C, the same for Sh. celti; D, SIPH of Sh. hangzhouensis; E, SIPH of Sh. celticola;
f, SIPH of Sh. szelegiewiczi (after Quednau, 1979).

- (Alata from leaf-nest.) BL more than 4mm. Hind wing with 2 oblique
veins arising from a common base. Secondary rhinaria on ANT III-V (not
VI), narrowly transverse but not extending more than half-way around
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
4 BL more than 4 mm. Rostrum longer than body ... Stomaphis yanonis
- BL less than 3 mm. Rostrum shorter than body . . . . . . . . . . . . . . . . . . 5
5 Antenna 4-segmented in aptera, 5-segmented in alata. Posterior abdomen
with lateral processes. SIPH tubular, swollen, constricted apically, and
with a subapical ring of 4 hairs (Fig. 38A) . . . . . . . . Sumatraphis celti
- Antenna 6-segmented. Abdomen without lateral processes. SIPH are
pores, very small cones, or truncate, without hairs . . . . . . . . . . . . . . . 6
130 Celtis

6 SIPH truncate, as long as ANT I (Fig. 38D). ANT PT/BASE 0.6-0.8. Rs

of forewing indistinct or absent . . . . . . . . . . . . Shivaphis hangzhouensis
- SIPH as very small cones, much shorter than ANT I, or merely pores.
ANT PT/BASE 0.15-0.5, Rs present at least distally . . . . . . . . . . . . . 7
7 ANT PT very short, about 3 times as long as wide or less . . . . . . . . 8
- ANT PT short, but at least 6 times longer than its maximum width
............................................................ 9
8 Apical part of CAUDA a rounded knob (Fig. 38B). ANT III of alata with
rhinaria in a line extending over most of length of segment . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shivaphis catalpinari
- Apical part of CAUDA finger-like, rather irregular in shape (Fig. 38C).
Alata with secondary rhinaria concentrated on central part of ANT III
.................................................. Shivaphis celti
9 SIPH very short cones, of height about half their basal width or less
(Fig. 38E). ANT PT/BASE about 0.3, PT about 0.6 of length of HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shivaphis celticola
- SIPH merely pores (Fig. 38F). ANT PT/BASE at least 0.4, and PT about
equal in length to HT II . . . . . . . . . . . . . . . . . . . . Shivaphis szelegiewiczi
10 CAUDA and SIPH dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- CAUDA pale, SIPH at least basally pale . . . . . . . . . . . . . . . . . . . . . . . 12
11 ANT IV of alata with 3-4 secondary rhinaria . . . . . . . Toxoptera celtis
- ANT IV of alata with 0-1 secondary rhinaria ..... Toxoptera aurantii
12 Head, legs and antennae of aptera mainly dark, femora basally pale but
with distal 0.6-0.7 black. SIPH slightly swollen over distal 0.7. CAUDA
with a constriction . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
- Head, legs and antennae of aptera mainly pale. SIPH not swollen.
CAUDA without a constriction . . . . . . . . . . . . . . . . Aulacorthum solani

CERATONIA Leguminosae

Ceratonia siliqua Aphis craccivora

(Carob)
 Chamaecyparis 131

CERCIDIPHYLLUM Cercidiphyllaceae

Cercidiphyllum japonicum Aulacorthum cercidiphylli

Myzus persicae
Key to aphids on Cercidiphylum
Inner faces of antennal tubercles parallel or slightly divergent in dorsal
view. ANT III of aptera usually with 1-3 small rhinaria near base. Femora
usually dark apically. Alata without a black dorsal abdominal patch
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum cercidiphylli
Inner faces of antennal tubercles convergent in dorsal view. ANT III of
aptera never with rhinaria. Alata with a black dorsal abdominal patch
................................................. Myzus persicae

CERCIS Leguminosae

Cercis canadensis Longistigma caryae

Aphis pawneepae
C. siliquastrum Myzus persicae
(Judas Tree)

CHAMAECYPARIS Cupressaceae

Chamaecyparis lawsoniana Cinara (Cedrobium) laportei

Cinara (Cupressobium)
louisianensis, tujafilina
Illinoia morrisoni
Key to aphids on Chamaecyparis
1 ANT PT/BASE more than 1. SIPH long and tubular, swollen distally
(Fig. 53A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni
- ANT PT/BASE less than 1. SIPH are broad, hairy cones ......... 2
2 Antenna 5-segmented. Dorsal hairs of aptera club-shaped, ornamented
with numerous barbules (Fig. 37A) ..... Cinara (Cedrobium) laportei
Antenna 6-segmented. Dorsal hairs normal, pointed . . . . . . . . . . . . . . 3
132 Chorisia

3 BL often more than 2.4mm. Aptera with dark sclerites on thorax and
ABD TERG 8. Apices of tibiae dark. ANT VI BASE with 8-14 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara (Cupressobiurri) tujafilina
BL less than 2.4 mm. Aptera without dark dorsal sclerites. Apices of tibiae
pale or dusky. ANT VI BASE with 5-6 hairs . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara (Cupressobium) louisianensis
N.B. Other aphids whose more normal hosts are Cedrus and Cupressus may
occasionally colonize Chamaecyparis, so it may be advisable to look also at
the keys to aphids on those plant genera.

CHORISIA Bombacaceae

Chorisia speciosa Myzus persicae

(Paneira)

CHOSENIA =SALIX

CHRYSOBALANUS Chrysobalanaceae

Chrysobalanus oblongifolius Aphis gossypii

CINCHONA Rubiaceae

Cinchona ledgeriana Aphis gossypii, spiraecola

(Quinine) Toxoptera aurantii, odinae
(Use key to polyphagous aphids, p. 532)
 Cinnamomum 133

CINNAMOMUM Cinnamons Lauraceae

B & E (1984) provided keys to the aphids recorded from camphor

(C. camphora) and cinnamon (C. zeylanicum). Here we treat the genus
Cinnamomum as a whole.

Host Plant List

Cinnamomum brevifolium Euthoracaphis umbellulariae
C. burmanni Sinomegoura citricola
C. camphora Aiceona actinodaphnis, japonica
(Camphor Tree) Euthoracaphis umbellulariae
Machilus machili
Sinomegoura citricola
Toxoptera aurantii, citricola
C. iners Euthoracaphis heterotricha
C. japonicum Euthoracaphis umbellulariae
Sinomegoura citricola
Thoracaphis sp.
C. pedunculatum Euthoracaphis umbellulariae
C. zeylanicum Euthoracaphis heterotricha
(Cinnamon Tree) [Micromyzus nigrum van der Goot,
1918]
Sinomegoura citricola

Key to aphids on Cinnamomum

1 SIPH longer than basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
SIPH either inconspicuous pores or broad cones, shorter than basal width
............................................................. 4
2 Body spindle-shaped. CAUDA very large with pointed apex. SIPH dark
apically, pale on basal half (Fig. 121B) . . . . . . . . Sinomegoura citricola
Body broadly oval. CAUDA rounded at apex. SIPH wholly dark ... 3
3 CAUDA with usually more than 25 hairs. Longest hairs on ANT III
longer than basal diameter of segment. BL often 2 mm or more .....
............................................. Toxoptera citricidus
CAUDA with usually less than 20 hairs. Longest hairs on ANT III shorter
than basal diameter of segment. BL less than 2 mm . . . . . . . . . . . . . . . .
.............................................. Toxoptera aurantii
134 Cinnamomum

Fig. 39. A, dorsal view of body of a Euthoracaphis; B, prosomal hairs of Eu. heterotricha; C, prosomal
hairs of Eu. umbellulariae.

4 Body of aptera aleyrodiform; subcircular, sclerotized, with much reduced

legs and antennae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Body of aptera of normal aphid type with well-developed legs and
antennae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5 Dorsal area of prosoma entire . . . . . . . . . . . . . . . . . . . . Thoracaphis sp.
Dorsal area of prosoma with distinct sutures dividing it into a large
quadrate central area, plus an anterior and two dorsolateral areas
(Fig. 39A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Prosoma with two longitudinal spinal rows of long thick hairs (80-125 µm
long) in addition to many long and short fine hairs (Fig. 39B) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euthoracaphis heterotricha
- Prosoma with all hairs fine, the longest of them 40-80 µm long (Fig. 39C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euthoracaphis umbellulariae
7 SIPH in form of small sclerotized rings. Wax glands evident. Hairs rather
sparse and mainly short . . . . . . . . . . . . . . . . . . . . . . Machilaphis machili
SIPH in form of broad hairy cones. Wax glands not evident. Hairs long
and numerous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 R IV+V shorter than HT II . . . . . . . . . . . . . . . . . . . . . Aiceona japonica
R IV+V longer than HT II . . . . . . . . . . . . . . . . . . Aiceona actinodaphne

CITHAREXYLUM Verbenaceae

Citharexylum fruticosum Aphis spiraecola

 Combretum 135

C. myrianthum Myzus persicae

(Use key to polyphagous aphids, p. 532)

CITRUS Rutaceae

See B & E (1984) for a key to aphids on Citrus.

COCOS Palmae

Cocos nucifera Astegopteryx nipae, rappardi,

rhapidis
Cerataphis lataniae
(For palm aphids see B & E, 1984)

COMBRETUM Combretaceae

Apart from a few polyphagous species, the aphid fauna of Combretaceae and
Burseraceae consists entirely of members of the genus Paoliella. Species
of Paoliella are specific at least at the level of host genera (Combretum,
Commiphora, Terminalia), with eight known from Combretum. Three of
these are only known as alate morphs, and one only from a single aptera, so
apterae and alatae are keyed separately.

Host Plant List

Combretum angolense Paoliella chiangae
C. apiculatum Paoliella nachensis, papillata
C. ghansalense Aphis gossypii
C. glomeruliflorum Myzus persicae
C. gueinzii = C. molle
C. krausii Aphis gossypii, spiraecola
C. mechowianum Aphis gossypii
C. molle Aphis gossypii
136 Combretum

Paoliella browni, namaachae,

papillata, wettsteini
C. psidioides Aphis gossypii
Paoliella browni ssp. capinganae,
chiangae, monotuberculata,
nachensis, papillata
C. zeyheri Paoliella papillata
Combretum sp. Paoliella longirostris
Toxoptera aurantii

Key to aphids on Combretum

l SIPH conspicuous, longer than their basal width. Dorsal processes absent.
CAUDA tongue-shaped or finger-like (sometimes with a mid-way con-
striction), without a knobbed apex. Anal plate entire . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
(Aphis gossypii, A. spiraecola, Toxoptera aurantii, Myzus persicae)
SIPH not longer than basal width, inconspicuous except when pigmented.
CAUDA with an apical knob, anal plate bilobed. Often with conspicuous
dorsal processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Apterous viviparous females (and intermediates) . . . . . . . . . . . . . . . . . . 3
Alate viviparous females ....................................... 5
3 Dorsal processes maximally about 4 x their middle diameter, cylindrical
or club-shaped, with apex about as wide as base (Fig. 40A) . . . . . . . . .
.............................................. Paoliella papillata
Dorsal processes very much longer, broad at base and tapering to much
narrower apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 ABD TERG 8 with two separate short finger-like processes. Knob of
CAUDA clearly longer than its greatest width. First tarsal segments
usually with 5-6 ventral hairs . . . . . . . . . . . . . . . . . . . Paoliella wettsteini
ABD TERG 8 with a median shallow protrusion bearing two hairs. Knob
of CAUDA as wide as long. First tarsal segments usually with 2-3 ventral
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella browni
5 Tip of forewing with a large dark spot, in addition to the dark spot at
the end of Culb. ANT PT/BASE more than 2.5 . . . . . . . . . . . . . . . . . 6
Tip of forewing without a dark spot, although such a spot often present
at the end of Culb. ANT PT/BASE less than 2.0 . . . . . . . . . . . . . . . . 7
6 R IV+V shorter than HT II . . . . . . . . . . . . . . . . . . . . Paoliella nachensis
- R IV+V longer than HT II . . . . . . . . . . . . . . . . . . . Paoliella namaachae
7 Fore femora similar in size and thickness to mid- and hind femora.
Antenna 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
 Combretum 137

Fig. 40. Typical spinal processes of Paoliella spp.: A, papillata (aptera), B, papillata (alata),
C, chiangae (alata), D, longirostris (alata), E, delottoi (aptera or alata), F, commiphorae (alata),
G, ufuasi (aptera), H, hystrix (aptera), I, echinata (aptera), J, ayari (aptera), K, mesothoracic process
of ayari (alata).

- Fore femora much enlarged in comparison to mid- and hind femora.

Antenna 6-segmented (antenna of P. longirostris not seen) ........ 9
8 ANT PT/BASE 0.54-0.73. R IV+V 0.70-0.82 x HT II . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella monotuberculata
- ANT PT/BASE 0.31-0.55. R IV+V 0.82-0.96 x HT II . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella browni capinganae
9 Head, pterothorax, siphunculi and dorsal and lateral processes all pale.
No pigmented dorsal abdominal sclerites . . . . . . . . . Paoliella wettsteini
- Head, pterothorax, siphunculi and at least the bases of the lateral
processes dusky to dark. Dark dorsal abdominal sclerites often present
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Dorsal processes almost cylindrical for most of length, or club-shaped,
with broadly rounded apices (Fig. 40B). R IV+V a little longer than
(1.05-1.15 x) HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella papillata
138 Commiphora

Dorsal processes tapering, conical or finger-like. R IV+V either shorter

than or much longer than HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Femora dark. R IV+V shorter than HT II. Spinal processes pigmented;
those on ABD TERG 1 less than 3 x their basal widths (Fig. 40C), and
at least those on ABD TERG 3 and 4 have transverse sclerotic bars uniting
their bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella chiangae
Femora pale. R IV+V at least twice as long as HT II. Spinal processes
pale, with basal sclerites pale or only weakly pigmented; spinal processes
on ABD TERG 1 about 4 x their basal widths (Fig. 40D) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella longirostris

COMMIPHORA Burseraceae

This genus has a similar aphid fauna to Combretum, q.v., but with different
species of Paoliella.

Host Plant List

Commiphora africana Aphis gossypii
Paoliella delottoi
C. dulcis Paoliella delottoi
C. fischeri Paoliella commiphorae ssp.
persimilis
C. mollis Toxoptera citricidus
C. neglecta Toxoptera aurantii
C. pedunculata Paoliella ayari
C. pilosa Paoliella echinata, ufuasi
C. savojae Paoliella hystrix
C. zimmermani Paoliella commiphorae

Key to aphids on Commiphora

1 SIPH conspicuous, longer than their basal width. Dorsal processes absent.
CAUDA tongue-shaped, anal plate entire . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
(Aphis gossypii, Toxoptera aurantii, Toxoptera citricidus)
SIPH not longer than basal width, inconspicuous except when pigmented.
CAUDA with apical knob, anal plate bilobed. Often with conspicuous
dorsal processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
 Commiphora 139

2 Apterous viviparous females (and intermediates) . . . . . . . . . . . . . . . . . . 3

Alate viviparous females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3 Dorsal processes maximally about 1.5 x their basal widths, with bluntly
rounded apices (Fig. 40E). ANT PT/BASE 0.4-0.5. Fore tibiae often with
rounded scent plaques . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella delottoi
Dorsal processes long and tapering, the longest at least 3 x their basal
widths. ANT PT/BASE more than 0.5. Fore tibiae without scent plaques
............................................................. 4
4 Dorsal processes tapering towards their rounded or pointed apices,
without a trace of apical swelling (Figs 40G, H) . . . . . . . . . . . . . . . . . 5
Dorsal processes cylindrical or more-or-less expanded at their apices
(Figs 40I, J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 Pleural processes absent or much reduced; most segments with only spinal
and lateral processes, which are maximally about as long as ANT III
(Fig. 40G). R IV+V about equal in length to HT II . . . . . . . . . . . . . . . .
................................................. Paoliella ufuasi
Pleural processes present on anterior abdominal segments, and of similar
length to spinal and lateral processes, which are maximally more than
2 x ANT III (Fig. 40H). R IV+V 1.1-1.2 X HT II . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella hystrix
6 ANT PT/BASE 0.8-0.9. Dorsal processes strongly flared at apices
(Fig. 40J), the longest less than 0.3 x BL . . . . . . . . . . . . Paoliella ayari
ANT PT/BASE 0.5-0.6. Dorsal processes only weakly expanded or
cylindrical at apices (Fig. 40I), the longest of them more than 0.3 x BL
............................................... Paoliella echinata
7 ANT PT/BASE about 0.5 or less. Fore femora not clearly thicker than
mid- or hind femora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
ANT PT/BASE at least 0.7 Fore femora enlarged in comparison with
mid- and hind femora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Head and thorax without processes. Dorsal abdominal processes short and
stout, the longest less than 1.5 x their basal widths (Fig. 40E). ANT HI
with 9-16 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella delottoi
Head with tubercles and thorax with finger-like processes. Dorsal
abdominal processes finger-like, the longest 2 or more x their basal
width. ANT III with 18-32 rhinaria . . . . . . . . . . . . . . Paoliella echinata
9 ANT PT/BASE more than 1.5. Dorsal processes all tubercular, less than
2 x their basal widths (Fig. 40F). Radial sector and media of forewing
bordered with fuscous distally . . . . . . . . . . . . . . Paoliella commiphorae
- ANT PT/BASE less than 1.5. Dorsal processes on thorax and ABD TERG
140 Conopharyngia

1-3 finger-like, the longest at least 2 x their basal widths. Radial sector
and media not bordered distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Mesothoracic dorsal processes dark and heavily spiculose (Fig. 40K).
Forewing vein Cu1b, bordered with fuscous . . . . . . . . . . . Paoliella ayari
Mesothoracic dorsal processes pale and no more spiculose than those on
other tergites. Forewing vein Culb not bordered ..... Paoliella ufuasi

CONOPHARYNGIA = TABERNAEMONTANA

CORCHORUS Tiliaceae

Corchorus capsularis Uroleucon compositae

CORDIA Boraginaceae

Host Plant List

Cordia abyssinica Myzus persicae
C. alliodora Aphis spiraecola
Toxoptera aurantii
C. corymbosa Uroleucon ambrosiae
C. myxa Aphis gossypii
C. nitida Aphis spiraecola
(Use key to polyphagous aphids, p. 532)

CORNUS Dogwoods Cornaceae

Cornus is the primary host of the genus Anoecia, and the primary or only host
of seven species of Aphis. In both these genera some species are difficult to
separate, and identifications made with the following key need to be treated
 Cornus 141

with caution. The North American Cornus-feeding species of the genus Aphis
were reviewed by Robinson and Chen (1969). For Anoecia only the alatae (i.e.
the spring migrant fundatrigeniae leaving Cornus in spring, and the sexuparae
returning to it in autumn) are keyed, as in several species the only apterous
morphs on Cornus are the fundatrix in spring and the sexuales in autumn.

Host Plant List

Cornus alba Anoecia corni, oenotherae
Aphis salicariae
C. alternifolia Macrosiphum hamiltoni
C. amomum Anoecia cornicola, oenotherae
Aphis cornifoliae, spiraecola
Myzus persicae
C. brachypoda Anoecia fulviabdominalis
Sitobion cornifoliae
C. capitata Myzus ornatus
C. chinensis Myzus persicae
C. controversa Anoecia fulviabdominalis, Anoecia
sp.
Toxoptera odinae
C. coreana Anoecia fulviabdominalis
Toxoptera odinae
C. florida Aphis cornifoliae
C. kousa Sitobion cornifoliae
C. mas Anoecia sp.
Aphis gossypii, helianthi
Macrosiphum euphorbiae
C. microcarpa Pseudasiphonaphis corni
C. paniculata Anoecia oenotherae
Aphis cornifoliae, [maculatae?]
C. racemosa Aphis caliginosa
(Grey Dogwood) Myzus persicae
C. sanguinea Anoecia corni, major, vagans
(Bloodtwig Dogwood) Aphis cornifoliae, fabae,
helianthi, salicariae, spiraecola
C. stolonifera Anoecia cornicola, oenotherae
(Red-Osier Dogwood) Aphis cornifoliae, helianthi,
neogillettei, nigratibialis,
salicariae, spiraecola
142 Cornus

Fig. 41. SIPH and marginal tubercles on ABD TERG 5-7 of apterae of A, Anoecia corni alate sexupara,
B, Pseudosiphonaphis corni aptera and C, Aphis caliginosa aptera; SIPH of D, Aphis cornifoliae,
E, A. neogillettei and F, A. salicariae.

Aulacorthum solani
Macrosiphum euphorbiae,
hamiltoni, manitobensis
Myzus persicae
Cornus sp. Prociphilus cornifoliae

Key to aphids on Cornus (cannot be applied to fundatrices or sexual morphs)

1 ANT PT/BASE less than 0.5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE 1.0 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
2 Well-developed wax pore plates present on head, thorax and abdomen.
SIPH pores absent. BL of adult aptera more than 3 mm, and dorsum pale.
Forewing of alata with unbranched media and pterostigma much longer
than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus cornifoliae
- Wax pore plates not developed. SIPH present as pores on low hairy cones
(Fig. 41 A). BL of adult aptera less than 3 mm, and dorsum dark or at least
with dark cross-bands. Forewing of alata with once-branched media and
short, wide, black pterostigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Large, flat, round marginal tubercles situated near spiracles on ABD
TERG 1-7, including 5 and 6 (anterioventrad and posterioventrad to
 Cornus 143

SIPH, dorsad to spiracles; arrowed in Fig. 41A), although sometimes

reduced in size, and occasionally missing on one or more segments
............................................................. 4
- Large, flat, round marginal tubercles on ABD TERG 1-4 and 7, but
usually absent from 5 and always absent from 6 . . . . . . . . . . . . . . . . . 7
4 Alata without a large dark dorsal abdominal patch . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . spring, migrant of Anoecia vagans
- Alata with a large dark dorsal abdominal patch . . . . . . . . . . . . . . . . . . 5
5 Alata usually without secondary rhinaria on ANT V, and never on VI.
ABD TERG 3-5 often with one or more short spatulate hairs . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sexupara of Anoecia vagans
- Alata with 1-4 secondary rhinaria on ANT V, and 0-2 on ANT VI. ABD
TERG 3-5 with only acute hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 ANT III with 9-17 rhinaria. R IV+V usually less than 0.9 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoecia corni
- ANT III with 13-22 rhinaria. R IV+V usually more than 0.9 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoecia major
7 Alata with 0-3, often small, secondary rhinaria on ANT III, and without
a solid dark dorsal abdominal patch (but often in autumn with cross-
bands on ABD TERG 4-6) . . . . . . . . . . . . . . . . . . . . Anoecia oenotherae
- Alata with 4 or more, usually large, secondary rhinaria on ANT III, and
with a solid dark patch on ABD TERG 4-6 (except spring migrant of
A. cornicola?) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Alata with all hairs on ABD TERG 1-3 small, similar in size to those on
ABD TERG 4-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anoecia cornicola
- Alata with hairs on ABD TERG 1-3 mostly long and fine, more than twice
as long as those on ABD TERG 4-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Alata with HT II 0.55-0.63 x ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . spring migrant of Anoecia corni
- Alata with HT II 0.41-0.54 x ANT III .... Anoecia fulviabdominalis
10 SIPH very small, papillate. Marginal tubercles well developed, longer than
wide, of similar size on all of ABD TERG 1-7 (Fig. 41B) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudasiphonaphis corni
- SIPH tubular, tapering or cylindrical. Marginal tubercles absent or weakly
developed on ABD TERG 2-6; if present then smaller than on prothorax
and ABD TERG 1 and 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Antennal tubercles weakly developed or undeveloped . . . . . . . . . . . . 12
- Antennal tubercles well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
144 Cornus

12 Hairs on ANT III more than 2 x longer than basal diameter of segment.
SIPH clearly shorter than CAUDA. Closely-spaced spinulose ridges
present on ABD sternites 5 and 6 . . . . . . . . . . . . . . . . Toxoptera odinae
- Hairs on ANT III less than 2 x longer than basal diameter of segment.
SIPH usually as long as or longer than CAUDA. No evident spinulose
ridges on ABD sternites 5 and 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Hairs on ANT III short, maximally 35 µm, shorter than or about equal
to middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Hairs on ANT III long, maximally 35-75 µm, all or most clearly exceeding
middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
14 Marginal tubercles present on prothorax and on ABD TERG 2-5 as well
as 1 and 7; those on prothorax, 1 and 7 being exceptionally large and
broad-based (Fig. 41C) . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis caliginosa
- Marginal tubercles usually present only on prothorax and ABD TERG 1
and 7, and these not unusually large . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 CAUDA finger-like, with a slight constriction basad of midpoint, the
distal part elongate oval (Fig. 121T) . . . . . . . . . . . . . . . Aphis spiraecola
- CAUDA tongue-shaped or bluntly triangular, without elongate oval distal
part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Aptera with ANT III usually more than 1.3 x PT (except in very small
specimens). Alata with 15-36 secondary rhinaria on ANT III, and usually
none (sometimes 1-2) on ANT IV . . . . . . . . . . . . . . . . . . Aphis helianthi
- Aptera with ANT III usually less than 1.3 x PT. Alata with 3-16
secondary rhinaria on ANT III, and 0-5 on ANT IV . . . . . . . . . . . . 17
17 Hind femora, SIPH and CAUDA all uniformly dark. SIPH cylindrical,
only a little narrower at apex than at base, and often curved outwards
(Fig. 41D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis cornifoliae
- Hind femora pale (aptera) or pale on basal third (alata); SIPH dark,
CAUDA pale or dusky. SIPH tapering, much narrower at apex than at
base (especially in aptera), and usually rather straight . Aphis gossypii
18 Tibiae of cleared specimens uniformly brown-black .................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis nigratibialis
- Tibiae of cleared specimens pale, or dark only apically . . . . . . . . . . 19
19 SIPH long, about 0.25 of BL. Aptera with 15-22 secondary rhinaria on
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis maculatae
- SIPH much less than 0.25 of BL. ANT III of aptera usually without
secondary rhinaria (rarely 1-4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 CAUDA with 11-27 hairs. Dorsal hairs on ABD TERG 1-6 usually less
than 40 µm long. SIPH usually rather straight . . . . . . . . . . Aphis fabae
 Cornus 145

- CAUDA with 6-11 hairs. Dorsal hairs on ABD TERG 1-6 usually long,
more than 40µm. SIPH often curved outwards distally (Figs 41E, F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 ABD TERG 8 with 3-6 hairs. SIPH of aptera about 5 x longer than their
width at base (Fig. 41E). Alata without dark dorsal markings on ABD
TERG 1-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis neogillettei
- ABD TERG 8 with 6-12 hairs. SIPH of aptera only 2-3 x longer than
their width at base (Fig. 41F). Alata with dark bars or sclerotic patches
on ABD TERG 1-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis salicariae
22 Antennal tubercles with inner faces divergent in dorsal view (Fig. 42K).
SIPH with a subapical zone of polygonal reticulation . . . . . . . . . . . . 23
- Antennal tubercles with inner faces parallel or convergent in dorsal view
(Figs 42I, J). SIPH without subapical polygonal reticulation . . . . . . . 26
23 CAUDA long and finger-like, 0.12-0.20 of BL. Hairs on ANT III more
than 0.5 of middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . 24
- CAUDA short, less than 0.11 of BL. Hairs on ANT III less than 0.5 of
middle diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
24 SIPH 1.5-2.1 x CAUDA and reticulated over distal 13-25%. R IV+V
0.9-1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH 2.3-2.5 x CAUDA and reticulated over distal 6-11%. R IV+V
1.05-1.20 x HT II . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum hamiltoni
25 ANT BASE VI less than 0.2 mm, much shorter than ANT I + II measured
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum manitobensis
- ANT BASE VI about 0.3 mm, as long as ANT I + II measured together
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion cornifoliae
26 Antennal tubercles with inner faces approximately parallel in dorsal view
(Fig. 42J). ANT III of aptera usually with a single rhinarium near base.
SIPH straight, tapering from base to flange (Fig. 121K) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- Antennal tubercles with inner faces convergent in dorsal view (Fig. 42I).
ANT III of aptera never with any rhinaria. SIPH tapering with a slight
S-curve, or slightly swollen on distal half . . . . . . . . . . . . . . . . . . . . . . . 27
27 ANT PT/BASE less than 2.5. Aptera with a dorsal intersegmental pattern
of dark ornamentation (Fig. 121J). SIPH coarsely imbricated, with a
slight S-curve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus ornatus
- ANT PT/BASE more than 3.0. Aptera without any clear dorsal markings.
SIPH slightly swollen on distal half (Fig. 121H) . . . . . . Myzus persicae
146 Corylopsis

CORYLOPSIS Hamamelidaceae

Corylopsis gotoana Hydronaphis impatiens

C. pauciflora Hydronaphis impatiens
C. spicata Hydronaphis impatiens

CORYLUS Cobnuts and Hazelnuts Corylaceae

This is a revised version of the key given in B & E (1984).

Host Plant List

Corylus americana Macrosiphum (Neocorylobium)
coryli, pseudocoryli
C. avellana Corylobium avellanae
(Common Filbert or Hazelnut) Myzocallis coryli
Pterocallis (Paratinocallis)
corylicola
C. colurna Myzocallis coryli
Pterocallis affinis
C. cornuta (incl. var. californica) Corylobium avellanae
[Illinoia corylina]
Macrosiphum (Neocorylobium)
coryli, pseudocoryli,
vandenboschi
C. heterophylla (incl. var. thunbergii) Macrosiphum (Neocorylobium)
corylicola
Neochromaphis coryli
Mesocallis pteleae
Pterocallis heterophyllus
Pterocallis (Paratinocallis)
corylicola
Tinocallis nikkoensis
C. mandshurica Betacallis alnicolens
Pterocallis (Paratinocallis)
corylicola
 Corylus 147

C. maxima Corylobium avellanae

Myzocallis coryli
C. sieboldiana (incl. var. brevirostris) Macrosiphum (Neocorylobium)
corylicola
Myzocallis coryli
Neochromaphis coryli
Pterocallis (Paratinocallis)
corylicola
Pterocallis montana
Tinocallis nikkoensis

Key to the aphids on Corylus

1 Head with a transverse black band ventrally between the eyes . . . . . . .
............................................. Betacallis alnicolens
- Head without a transverse ventral black band . . . . . . . . . . . . . . . . . . . . 2
2 CAUDA knobbed, anal plate bilobed. SIPH as small, truncate cones.
(Adult viviparae usually or always alate) . . . . . . . . . . . . . . . . . . . . . . . . 3
- CAUDA finger-shaped, anal plate entire. SIPH long and tubular, with a
subapical zone of polygonal reticulation. (Adult viviparae in colonies
usually mainly apterous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
3 ANT PT/BASE more than 2.0 . . . . . . . . . . . . . . . . . . . Myzocallis coryli
- ANT PT/BASE 0.1-1.2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Forewings with extensive pattern of dark markings. PT extremely short,
ANT PT/BASE only about 0.1 . . . . . . . . . . . . . . Neochromaphis coryli
- Forewings sometimes with pigment along veins and/or on pterostigma,
but without extensive pigmented areas. ANT PT/BASE at least 0.2
............................................................. 5
5 Spinal hairs on ABD TERG 1-4 long, up to 6 x basal diameter of ANT
III, pointed and spine-like, on low tuberculate bases . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis heterophyllus
- Spinal hairs on ABD TERG 1-4 all short, not or hardly exceeding basal
diameter of ANT III, or if longer then capitate . . . . . . . . . . . . . . . . . . 6
6 ANT PT/BASE 0.2-0.45 . . . . . . . . . . . . . . . . . . . . . . Pterocallis montana
- ANT PT/BASE 0.5 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Mesonotum and ABD TERG 2 and 3 each with a pair of large spinal hair-
bearing processes. ANT III with 15-19 secondary rhinaria . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis nikkoensis
- Mesonotum and ABD TERG 2 and 3 without spinal processes. ANT III
with 2-14 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
148 Crataegus

8 ANT III dark, with 7-14 rhinaria. Abdominal marginal tubercles each
bearing a single small hair . . . . . . . . . . . . . . . . . . . . . . Mesocallis pteleae
- ANT III pale or only dark at apex, with 2-11 rhinaria. Abdominal
marginal tubercles mostly with 2-3 hairs . . . . . . . . . . . . . . . . . . . . . . . . 9
9 ANT PT/BASE 0.5-0.9. ABD TERG 1-7 with 6-10 spinopleural hairs,
the spinal hairs often grouped on paired tubercles . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis affinis
- ANT PT/BASE 0.9-1.2. ABD TERG 1-7 each with a single pair of spinal
hairs only, those on 3, 5 and 7 being displaced laterally . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis (Paratinocallis) corylicola
10 Dorsal hairs of aptera long, thick and slightly capitate, arising from large
tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Corylobium avellanae
- Dorsal hairs of aptera shorter, not capitate, not borne on tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Dorsal cuticle strongly sclerotized and very wrinkled (although quite pale)
. . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) vandenboschi
- Dorsal cuticle not sclerotic, smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Hind tibiae and SIPH of aptera wholly black, CAUDA dusky to dark
. . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobiuni) corylicola
- Hind tibiae of aptera with paler middle section; SIPH wholly black or pale
at bases, CAUDA pale to dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Aptera with abdominal cuticle usually pigmented marginally, and also
dorsally across ABD TERG 5-7. SIPH wholly black, ABD TERG 8
and CAUDA contrastingly pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) coryli
- Aptera with abdomen wholly pale. SIPH usually paler at base, CAUDA
pale or dusky . . . . . . . . . . Macrosiphum (Neocorylobium) pseudocoryli

CRATAEGUS Hawthorns, May Trees Rosaceae

Numerous aphids utilize species of Crataegus as their primary hosts, so that

the only morphs occurring are the fundatrix and her progeny in spring (often
causing leaf deformation or galling), and the return migrants and sexuales in
autumn. The key that follows the host list is applicable as indicated to the
spring generations of such species, but cannot be used for the autumn pre-
sexual and sexual generations. Couplets 17-26 are based on the intensive work
of Stroyan (1958) and the key to Dysaphis on Crataegus by Shaposhnikov
and Marolev (1979); the species-level taxonomy of the genus Dysaphis is
 Crataegus 149

particularly difficult, and a series of specimens from a population will need to

be examined before much reliance can be placed on any species determination.
This part of the key only applies to alate spring migrants, but details of the
appearance of the gall and the fundatrix in life (see section III), together with
the species of Crataegus colonized, may help to confirm the identification.

Host Plant List

Crataegus acclivis Rhopalosiphum insertum
C. altaica Dysaphis sorbiarum
C. azarolus Dysaphis crataegi
C. brainerdii Rhopalosiphum insertum
C. clara Hyalomyzus eriobotryae
C. coccinea Eriosoma lanigerum
Nearctaphis crataegifoliae
Ovatus crataegarius
Prociphilus (Neoparacletus)
corrugatans
Prociphilus (Stagona) pini
Rhopalosiphum ?insertum
C. crusgalli Eriosoma crataegi, lanigerum
Hyalomyzus sensoriatus
Nearctaphis clydesmithi
Prociphilus (Neoparacletus)
corrugatans
Rhopalosiphum insertum
Utamphorophora crataegi
C. cuneata Eriosoma lanigerum
Prociphilus (Stagona)
[crataegicola], pini
Rhopalosiphum ?insertum
C. curvisepala Dysaphis incognita
C. douglasii Aphis pomi
Fimbriaphis gentneri
Nearctaphis bakeri, sclerosa
C. hissarica Dysaphis sorbiarum
C. laevigata = C. oxyacantha
C. x lavallei Fimbriaphis gentneri
Utamphorophora crataegi
C. maximowiczii [Ovatus malisuctus]
[Prociphilus kuwanai]
150 Crataegus

C. mexicana = C. pubescens
C. mollis Rhopalosiphum insertum
C. monogyna Aphis pomi
Dysaphis angelicae, apiifolia ssp.
petroselini, crataegi, lauberti,
ranunculi
Fimbriaphis gentneri
Nearctaphis bakeri
Ovatus crataegarius
Prociphilus (Stagona) pini
Rhopalosiphum insertum
C. orientalis Dysaphis crataegi ssp. heraclei
C. oxyacantha Aphis fabae, pomi
Aulacorthum solani
Dysaphis crataegi
Fimbriaphis gentneri
Macrosiphum euphorbiae
Nearctaphis crataegifoliae
Ovatus crataegarius, insitus
Prociphilus (Stagona) pini
Rhopalosiphum insertum
C. phaenopyrum Rhopalosiphum insertum
C. pinnatifolia Ovatus crataegarius
C. praeformosa Hyalomyzus tissoti
C. x prunifolia Rhopalosiphum insertum
C. pubescens Muscaphis mexicana
Rhopalosiphum insertum
C. punctata Eriosoma lanigerum
Muscaphis stroyani
Nearctaphis crataegifoliae
C. rivularis Eriosoma lanigerum
Nearctaphis bakeri
C. songorica Dysaphis crataegi ssp. pallida
C. succulenta Nearctaphis crataegifoliae
C. tomentosa Eriosoma crataegi
Prociphilus (Neoparacletus)
corrugatans
C. uniflora Eriosoma lanigerum group
Hyalomyzus eriobotryae
 Crataegus 151

Schizoneurata tissoti
Utamphorophora crataegi
C. vicana Hyalomyzus tissoti
Crataegus sp. Dysaphis laserpitii
[Dysaphis ramani Das and
Raychaudhuri, 1983]
Muscaphis canadensis
Rhopalosiphum nigrum, rufulum

Key to aphids on Crataegus

1 ANT PT/BASE less than 0.5. Antenna of alata with narrow, transversely
elongate or annular secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 0.5. Antenna of alata with oval or circular
secondary rhinaria ............................................ 6
2 SIPH pores present, their rims partly sclerotized. Wax gland pore plates
usually consisting of a ring of large facets around an undivided or partially
divided central area (Fig. 47C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- SIPH absent, at least in apterae. Wax gland pore plates with numerous
small facets and no central area, or with a group of large facets including
a few small separate clear areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Aptera with greatest diameter of SIPH pore 0.08-0.16 mm. ANT III
usually more than 1.25 x more than ANT IV+V . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma crataegi
- Aptera with maximum diameter of SIPH pore less than 0.06 mm. ANT
III usually less than 1.25 x more than ANT IV+V . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanigerum
4 Wax pore plates (in immatures) comprising groups of 6-20 large facets
often including a few small clear areas. Alata (= sexupara) with media
of forewing once-branched (adult aptera unknown) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneurata tissoti
- Wax pore plates with more numerous small facets. Alata (= spring
migrant) with media of forewing unbranched . . . . . . . . . . . . . . . . . . . . 5
5 Fundatrix with R IV+V longer than ANT BASE V. Spring migrant alata
with pterostigma of normal length, extending about one third of distance
from base of Rs to tip of wing, and with 0-4 rhinaria on ANT V ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus (Neoparacletus) corrugatans
- Fundatrix with R IV+V shorter than ANT BASE V. Spring migrant
alata with pterostigma elongate distally, extending about one half of
distance from base of Rs to tip of wing, and with 3-11 rhinaria on
ANT V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus (Stagona) spp.
152 Crataegus

6 CAUDA usually short, helmet-shaped or triangular, shorter than or about

as long as its basal width in dorsal view; if tongue-shaped and up to
1.4 x its basal width then SIPH have rows of minute nodules or spinules
on closely-spaced imbrications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- CAUDA tongue- or finger-shaped, clearly longer than its basal width;
sometimes less than 1.4 x longer, but then SIPH do not have closely-
spaced rows of minute nodules/spinules . . . . . . . . . . . . . . . . . . . . . . . . 27
7 Dorsal cuticle of aptera (fundatrix) very rough with numerous small
denticulate or wart-like projections, and clothed with numerous long,
fine, curved hairs mostly over 60 µm long (Fig. 42A). Alata without a solid
black dorsal abdominal patch, and with SIPH more than 2 x CAUDA
............................................................. 8
- Dorsal cuticle of aptera (fundatrix or subsequent generation) smooth, with
fewer, shorter hairs (maximally 50 µm). Alata usually with a solid black
dorsal abdominal patch, or if without then SIPH less than 2 x CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
8 SIPH of aptera (fundatrix) 0.25-0.28 mm long, 1.6-2.0 x ANT V BASE.
Alata with 12-22 secondary rhinaria ANT I I I , 6-11 on IV, 2-5 on V, and
ANT PT/BASE 1.7-2.6 . . . . . . . . . . . . . . . . . . . . . . Muscaphis mexicana
- SIPH of fundatrix 0.33-0.45 mm long, 2.2-3.0 x ANT V BASE. Alata
with 25-50 secondary rhinaria on ANT I I I , 15-33 on IV, 9-20 on V, and
ANT PT/BASE 3.0-4.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 CAUDA of fundatrix bluntly triangular (Fig. 42B). SIPH of fundatrix
4-5 x its minimum width, and that of alata 5-6 x its minimum width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Muscaphis canadensis
- CAUDA of fundatrix helmet-shaped, with a pointed apex (Fig. 42C).
SIPH of fundatrix 7-8 x its minimum width, and that of alata
6.5-8.5 x its minimum width . . . . . . . . . . . . . . . . . . Muscaphis stroyani
10 SIPH with closely-spaced rows of imbrications bearing numerous small
nodules or spinules (e.g. Fig. 42M) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- SIPH normally imbricated, the imbrications sometimes spinulose but not
in regular closely-spaced rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
11 Dorsum of aptera with a solid black central sclerite covering ABD TERG
3-5, or 3-6, and transverse bars on more posterior tergites . . . . . . . 12
- Dorsum of aptera with separate transverse bars on ABD TERG 5-8,
and with only irregular broken sclerites on more anterior tergites . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
12 Tibiae black. Aptera with dorsal patch extending laterally to marginal
sclerites, at least on ABD TERG 4-5 . . . . . . . Nearctaphis clydesmithi
- Tibiae mainly pale with black apices. Aptera with dorsal patch not
extending laterally to marginal sclerites . . . . . . . . . . . . . . . . . . . . . . . . . 13
 Crataegus 153

Fig. 42. A, dorsal cuticle of fundatrix of Muscaphis stroyani; B, CAUDA of Muscaphis canadensis
(fundatrix); C, CAUDA of M. stroyani (fundatrix); D, CAUDA of Nearctaphis bakeri (aptera); E, CAUDA of
Dysaphis crateaegi; F, dorsal abdominal patch of Dysaphis crataegi kunzei; G, the same for D. crataegi;
H, comparison of positions of marginal tubercle on TERG 7 of Rhopalosiphum (above) and Aphis (below);
I, inner faces of antennal tubercles convergent; J, inner faces of antennal tubercles parallel; K, inner
faces of antennal tubercles diverging; L, SIPH of Rhopalosiphum rufulum spring migrant alata; M, SIPH
of Nearctaphis bakeri aptera; N, SIPH of Hyalomyzus eriobotryae aptera; 0, SIPH of H. tissoti aptera;
P, SIPH of Utamphorophora crataegi aptera; Q, SIPH of Fimbriaphis gentneri aptera.
154 Crataegus

13 Alata with a solid black patch on ABD TERG 3-5, partially fused with
bar on 6 . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis californica nigrescens
- Alata without a solid black dorsal patch, only with transverse bars or
dashes on ABD TERG 5-8 . . . . . . . . . . . . . . . . . . . Nearctaphis sclerosa
14 R IV+V 0.14-0.18 mm long, usually with 4 accessory hairs. Alata without
a solid black dorsal abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- R IV+V 0.10-0.13 mm long, usually with 2 accessory hairs. Alata with a
solid black patch covering ABD TERG 3-5, or 3-6 . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis bakeri
15 Head of aptera blackish-brown, and SIPH darker than CAUDA. Alata
with transverse dark bars on ABD TERG 6-8 or 7-8 . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis crataegifoliae occidentalis
- Head of aptera pale, and SIPH paler than CAUDA. Alata with a complete
transverse dark bar only on ABD TERG 8 . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis crataegifoliae crataegifoliae
16 Spinal tubercles usually present and well developed on head, prothorax
and all abdominal segments. SIPH of aptera pale or dusky, less than
0.14 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis sorbiarum
- Spinal tubercles only on head and ABD TERG 7-8, rarely on other
segments. SIPH dark, more than 0.14 mm long . . . . . . . . . . . . . . . . . 17
17 Spring migrant alata (henceforth 's.m.a.') with marginal tubercles on ABD
TERG 7 in most specimens, frequently on both sides. Hairs on ABD
TERG 8 usually 10-25 µm long . . . . . . . . Dysaphis apiifolia petroselini
- S.m.a. without marginal tubercles on ABD TERG 7 in most specimens,
and very rarely on both sides. Hairs on ABD TERG 8 30-70 µm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 S.m.a. with total number of secondary rhinaria on both ANT V combined
7-35, or if 5-6 then a transverse dark bar is present on ABD TERG 2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
- S.m.a. with usually only 0-5 secondary rhinaria on both ANT V
combined, rarely up to 6 or 7. ABD TERG 2 without a dark bar ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
19 ABD TERG 1 and 2 of s.m.a. without dark bars, at most with small
scattered circular sclerites . . . . . . . . . . . . . . . . . . . . . . Dysaphis angelicae
- ABD TERG 2, or both 1 and 2, of s.m.a. with a transverse dark bar,
sometimes partly broken into irregular sclerites . . . . . . . . . . . . . . . . . . 20
20 ANT III of s.m.a. with 44-75 secondary rhinaria, ANT IV with 15-35 and
ANT V with 3-19. R IV+V with 4-7 accessory hairs . . . . . . . . . . . . . . .
............................................. Dysaphis ranunculi
 Crataegus 155

- ANT III of s.m.a. with 37-47 secondary rhinaria, ANT IV with 10-19 and
ANT V with 2-6. R IV+ V with 2-3 accessory hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis virgata
21 ANT III of s.m.a. usually with 74-125 secondary rhinaria per mm of
length ...................................................... 22
- ANT III of s.m.a. usually with 130-192 secondary rhinaria per mm of
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22 R IV+V of s.m.a. 1.01-1.25 X HT II. Hairs on ANT III blunt or weakly
capitate, 9-21 µm long, those on ABD TERG 3 12-23 µm long . . . . . .
.............................................. Dysaphis incognita
- R IV+V of s.m.a. 0.83-0.98 x HT II. Hairs on ANT III mostly pointed,
20-29 µm long, those on ABD TERG 3 25-35 µm long . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis crataegi pallida
23 S.m.a. with hairs on ABD TERG 2 and 3 pointed, the longest 20-57 µm
long, but rarely less than 25 µm . . . . . . . . . . . . . . . . . . Dysaphis lauberti
- S.m.a. with hairs on ABD TERG 2 and 3 blunt, the longest 9-29 µm, but
rarely more than 25 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 S.m.a. with SIPH 2.9-3.5 x their middle diameter . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis crataegi heraclei
- S.m.a. with SIPH 3.6-5.7 x their middle diameter . . . . . . . . . . . . . . 25
25 S.m.a. with dorsal abdominal patch on ABD TERG 3-6 interrupted by
narrow gaps or lacunae between 3-4 and 4-5, as well as 5-6 (Fig. 42F).
Anterior part of subgenital plate with 4-10 hairs . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis crataegi kunzei
- S.m.a. with dorsal abdominal patch on ABD TERG 3-6 almost entire,
except for a narrow central gap between 5 and 6 (Fig. 42G). Anterior part
of subgenital plate with 2-7 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 (Requires large samples) Most s.m.a. with at least 1 antenna bearing 1 (or
more) secondary rhinaria on ANT V [mean number per antenna 0.53
(UK)-1.1 (former USSR)] . . . . . . . . . . . . . . . . Dysaphis crataegi crataegi
- Most s.m.a. without any secondary rhinaria on ANT V of either antenna
[mean number 0.32 per antenna (UK)] ... Dysaphis crataegi aethusae
27 Antennal tubercles undeveloped or weakly developed. SIPH of aptera
dark or, if only dark at apex, then short (less than 0.25 mm). CAUDA
and anal plate dusky to dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Antennal tubercles moderately to well developed. SIPH of aptera pale or
dusky, or dark only at apices and then over 0.3 mm long . . . . . . . . 34
28 SIPH slightly swollen or cylindrical on at least basal 0.7 of length, narrow-
ing only towards apex, and with a well-developed apical flange (e.g.
156 Crataegus

Fig. 42L). Marginal tubercles on ABD TERG 7 placed posteriodorsad

of spiracle, and usually smaller at base than the spiracular opening
(Fig. 42H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
- SIPH tapering from base to apex, and flange only moderately developed.
Marginal tubercles on ABD TERG 7 placed posterioventrad of spiracle
and usually larger than spiracular aperture . . . . . . . . . . . . . . . . . . . . . 31
29 Tibiae wholly dark . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum nigrum
- Tibiae mainly pale or dusky, sometimes dark at apices . . . . . . . . . . . 30
30 ANT PT/BASE less than 2.0 in fundatrix, and less than 2.5 in subsequent
generations (apterae and spring migrant alatae). SIPH of alata usually
less than 0.2 mm long (Fig. 42L) . . . . . . . . . . . Rhopalosiphum rufulum
- ANT PT/BASE more than 2.0 in fundatrix, and more than 2.5 in subse-
quent generations. SIPH of alata usually more than 0.2mm long . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum insertum
31 Aptera with transverse bars on ABD TERG 7 and 8 and at least some dark
spinopleural markings anterior to siphunculi. Alata with a series of dorsal
abdominal transverse bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis fabae
- Aptera without any dark dorsal markings, alata usually with only mar-
ginal abdominal sclerites anterior to siphunculi . . . . . . . . . . . . . . . . . . 31
32 CAUDA of aptera paler than SIPH, without any trace of a constriction,
and bearing 4-8 hairs (Fig. 121U) . . . . . . . . . . . . . . . . . . . Aphis gossypii
- CAUDA of aptera as dark as SIPH, with a slight constriction near its
midpoint, and bearing 7-19 hairs (Fig. 121T) . . . . . . . . . . . . . . . . . . . 33
33 Marginal tubercles present on ABD TERG 2-4. CAUDA with 10-19 hairs
(rarely less than 13). R IV+V more than 0.130mm . . . . . . Aphis pomi
- Marginal tubercles absent from ABD TERG 2-4. CAUDA with 7-15 hairs
(rarely more than 12). R IV+V less than 0.125 mm . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
34 Aptera with inner faces of antennal tubercles convergent in dorsal view
(e.g. Fig. 121I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
- Aptera with inner faces of antennal tubercles parallel or divergent in
dorsal view (e.g. Figs 121J, K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
35 SIPH slightly to moderately swollen over distal 0.5 to 0.7 of length. ANT
PT/BASE less than 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
- SIPH tapering gradually from base to flange. ANT PT/BASE more than
5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
36 ANT PT/BASE 2.5 or less . . . . . . . . . . . . . . . . Hyalomyzus sensoriatus
- ANT PT/BASE more than 3.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
 Craterispermum 157

37 R IV+V less than 1.1 x HTII. CAUDA with 6-11 hairs (usually 7). Alata
with a black dorsal abdominal patch . . . . . . . . . . . . . . . Myzus persicae
- R IV+V more than 1.1 x HT II. CAUDA with 4-6 hairs. Alata without
a black dorsal abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 SIPH of aptera moderately swollen, and only moderately imbricated
(Fig. 42N). ANT PT less than 1.1 x ANT III. Alata with 35 or more
secondary rhinaria on ANT III . . . . . . . . . . . . Hyalomyzus eriobotryae
- SIPH of aptera only slightly swollen, and coarsely imbricated (Fig. 42O).
ANT PT more than 1.2 x ANT III. Alata with 11-25 secondary rhinaria
on ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalomyzus tissoti
39 Spring migrant alata with 60-83 secondary rhinaria on ANT III, 36-52
on IV and 13-22 on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus insitus
- Spring migrant alata with 22-49 rhinaria on ANT III, 5-20 on IV and 0-10
on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus crategarius
40 SIPH with a subapical zone of polygonal reticulation (Fig. 121C). Hairs
on ANT III more than 0.5 x basal diameter of segment . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH without any polygonal reticulation. Hairs on ANT III less than
0.5 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41 SIPH swollen on about distal 0.5 of length (Fig. 42P). Alata without dark
dorsal abdominal markings . . . . . . . . . . . . . . . Utamphorophora crataegi
- SIPH tapering, cylindrical or only slightly swollen subapically (e.g.
Fig. 42Q). Alata with dark dorsal abdominal markings . . . . . . . . . . . 42
42 SIPH more than 2 x CAUDA, and in aptera often dark-tipped (Fig. 121K).
Dorsal abdomen of alata with transverse dark bars . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- SIPH less than 2 x CAUDA, not dark-tipped in aptera (Fig. 42Q). Dorsal
abdomen of alata with an almost solid dark patch across ABD TERG 4-5
or 3-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri

CRATERISPERMUM Rubiaceae

Craterispermum laurinum Aphis gossypii

Myzus persicae
Sitobion sp.
158 Cratoxylum

Key to aphids on Craterispermum

- SIPH with a subapical zone of polygonal reticulation ..... Sitobion sp.
- SIPH without polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

CRATOXYLUM Guttiferae

Cratoxylum celebicum Greenideoidea (Neogreenideoida)

philippensis

CROSSOPTERYX Rubiaceae

Crossopteryx febrifuga Aphis gossypii

Sitobion halli

Key to aphids on Crossopteryx

SIPH with a subapical zone of polygonal reticulation . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion halli
- SIPH without polygonal reticulation . . . . . . . . . . . . . . . . Aphis gossypii
(or try key to polyphagous aphids, p. 532)

CROTON Euphorbiaceae

Croton humulis Aphis spiraecola

C. macrostachys Eonaphis crotonis
Myzus persicae
C. subgratissimus Eonaphis crotonis
Croton sp. Toxoptera aurantii

Key to aphids on Croton

- Aptera and alata with very large spine-like processes, 4 per segment,
longer than the short, black, conical SIPH which have a subapical ring
of 4 hairs (Fig. 43) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eonaphis crotonis
 Cudrania 159

Fig. 43. Posteror abdominal segments of aptera of Eonaphis crotonis.

Aptera and alata without spine-like processes. SIPH tubular or tapering,

longer than basal width, without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . see key to polyphagous aphids, p. 532

CRYPTOMERIA Taxodiaceae

Cryptomeria japonica Cinara fresai

[Cinara greeni (Schouteden, 1905)]

CUDRANIA=MACLURA
160 X Cupressocyparis

X CUPRESSOCYPARIS Cupressaceae

X Cupressocyparis leylandii Illinoia morrisoni

CUPRESSUS Cypresses Cupressaceae

Cupressus has a small but diverse aphid fauna, with no evidence of host
specificity at the species level. The only possible confusion is between the
Cinara species, which are all in the subgenus Cupressobium.

Host Plant List

Cupressus arizonica Cinara cupressi, fresai
C. bakeri Siphonatrophia cupressi
C. benthami Cinara cupressi
Stomaphis cupressi
C. brunniana Cinara fresai
C. funebris Aphis craccivora
C. goveniana Cinara cupressi
C. guadalupenis Cinara fresai
Siphonatrophia cupressi
C. lambertiana Cinara cupressi
C. lusitanica Cinara cupressi
Siphonatrophia cupressi
C. macrocarpa Cinara cupressi, fresai
Illinoia morrisoni
Siphonatrophia cupressi
C. pygmea Cinara fresai
Siphonatrophia cupressi
C. sabiniana Cinara fresai
C. sempervirens Cinara cupressi, fresai
Siphonatrophia cupressi
Stomaphis cupressi ssp. caucasica
C. sargentii Siphonatrophia cupressi
 Cupressus 161

Fig. 44. A, SIPH of Siphonatrophia cupressi; B, ANT VI of Cinara cupressi; C, ANT VI of C. fresai.

C. torulosa Cinara cupressi, fresai

Cupressus sp. Cinara louisianensis, tujafilina

Key to aphids on Cupressus

1 ANT PT/BASE more than 0.5. CAUDA tongue- or finger-shaped, longer
than its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE less than 0.5. CAUDA broadly rounded . . . . . . . . . . 4
2 ANT PT/BASE about 1.0. SIPH pores on very small, shallow cones
(Fig. 44A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphonatrophia cupressi
- ANT PT/BASE more than 1.3. SIPH tubular, longer than basal width
............................................................. 3
3 Body spindle-shaped, without dark dorsal markings. SIPH long and
markedly swollen on distal half (Fig. 53A) . . . . . . . . Illinoia morrisoni
- Body oval, dorsal abdomen with an extensive black patch in aptera and
transverse bars in alata. SIPH tapering from base to flange . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
4 Rostrum much longer than body . . . . . . . . . . . . . . . Stomaphis cupressi
Rostrum much shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Femora pale, and tibiae wholly pale or dark only at apices (sometimes very
localized pigmentation at tibio-femoral joints) . . . . . . . . . . . . . . . . . . . . 6
Distal parts of femora and bases of tibiae dusky or dark . . . . . . . . . 7
6 BL often more than 2.4mm. Aptera with dark sclerites on thorax and
ABD TERG 8. Apices of tibiae dark. ANT VI BASE with 8-14 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara tujafilina
BL less than 2.4 mm. Aptera without dark dorsal sclerites. Apices of tibiae
pale or dusky. ANT VI BASE with 5-6 hairs ... Cinara louisianensis
162 Cussonia

7 Aptera with ANT VI BASE bearing 4-7 hairs confined to basal half
(Fig. 44B). R IV 0.12-0.17 mm long, bearing 2-4 accessory hairs; 1.4-1.9 x
HT I. Alata with 1-6 secondary rhinaria restricted to distal half of ANT
III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara cupressi
- Aptera with ANT VI BASE bearing 7-12 hairs, some on distal half
(Fig. 44C). R IV 0.16-0.24 mm long, bearing 5-7 accessory hairs; 1.8-
2.2 x HT I. Alata with 6-11 secondary rhinaria extending over most of
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara fresai

CUSSONIA Araliaceae

Cussonia natalensis Macrosiphum euphorbiae

C. paniculata Myzus persicae
Toxoptera odinae
C. spicata Aphis fabae, gossypii
Toxoptera odinae
C. umbellifera Macrosiphum euphorbiae
(Use key to polyphagous aphids, p. 532)

CYATHEA Tree Fern Cyatheaceae

Cyathea aquilina Aphis spiraecola

Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

CYCAS Cycadaceae

Aphis gossypii
 Dalbergia 163

CYNOMETRA Leguminosae

Cynometra cauliflora Aphis gossypii

Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

CYRILLA Cyrillaceae

Cyrilla racemiflora [Aphis nigra Wilson]

DAEMONOROPS Palmae

Daemonorops geniculata Astegopteryx nipae

(For palm aphids see B & E, 1984)

DALBERGIA Leguminosae

No aphids are recorded from commercially important rosewoods such as

D. nigra and D. stevensonii.

Dalbergia ferruginea Anomalosiphum sp.

D. hupeana Chuansicallis chengtuensis
Tinocallis (Quednaucallis)
nigropunctata
[Uroleucon compositae]
D. sissoo Acyrthosiphon pisum
Aphis craccivora, spiraecola
Myzus persicae
D. torta Anomalosiphum sp.
164 Dendrocalamus

Key to aphids on Dalbergia

1 SIPH with a subapical ring of hairs. Aptera with long thin hair-bearing
processes on ABD TERG 7 and 8 . . . . . . . . . . . . Anomalosiphum spp.
- SIPH without hairs. ABD TERG 7 and 8 with or without processes
............................................................. 2
2 CAUDA knobbed, anal plate bilobed. ANT PT/BASE a little less than
1 ........................ ................................... 3
- CAUDA tongue- or finger-shaped, anal plate entire. ANT PT/BASE
much more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 ABD TERG 1 and 2, and 7 and 8, bearing pale dorsal tubercles, with
traces of tubercles also on ABD TERG 3-6 . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis (Quednaucallis) nigropunctata
Dorsal abdominal tubercles not developed; ABD TERG 1 and 2 with
dusky spinal sclerites . . . . . . . . . . . . . . . . . . . . Chuansicallis chengtuensis
4 CAUDA and SIPH pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- CAUDA and SIPH black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 SIPH slightly swollen on distal half. Antennal tubercles scabrous, with
inner faces convergent in dorsal view . . . . . . . . . . . . . . . Myzus persicae
SIPH long and thin, tapering from base to flange. Antennal tubercles
smooth with inner faces divergent in dorsal view . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon pisum
6 Body spindle-shaped, CAUDA long and pointed with 11-18 hairs. SIPH
with an apical zone of polygonal reticulation . . . . . . . . . . . . . . . . . . . . . .
........................................... Uroleucon compositae
Body oval, CAUDA with 4-14 hairs. SIPH without polygonal reticulation
............................................................. 7
7 Aptera with an extensive black dorsal patch, alata with variably developed
transverse bars on ABD TERG 1-6 . . . . . . . . . . . . . . . Aphis craccivora
Neither aptera nor alata with dark dorsal markings . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola

DENDROCALAMUS Gramineae

Compared with the extensive aphid flora on Bambusa and Arundinaria,

few species are recorded from Dendrocalamus. Species within the genera
Astegopteryx and Pseudoregma cannot readily be differentiated, except for
P. alexanderi. Alternatively, try the key to aphids on Bambusa.
 Diospyros 165

Key to the aphids on Dendrocalamus

1 Adult viviparae all alate with long spinal and marginal hair-bearing finger-
like tubercles on abdomen (Fig. 30E) . . . . . . . . . Chucallis bambusicola
Adult viviparae apterae and/or alatae, without finger-like tubercles
............................................................. 2
2 Head of aptera without frontal horns. Dorsal and marginal body hairs
thick and spine-like . . . . . . . . . . . . . . . . . . . . . . . Glyphinaphis bambusae
Head of aptera with a pair of frontal horns. Body hairs short and/or
fine ......................................................... 3
3 CAUDA and anal plate rounded. Frontal horns of adult aptera usually
longer than ANT I and II combined . . . . . . Ceratoglyphina bambusae
CAUDA knobbed, anal plate bilobed. Frontal horns of adult aptera not
longer than ANT I and II combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 SIPH pores on shallow cones encircled by fine hairs. Aptera with wax
pore plates (when present) usually arranged in marginal longitudinal rows
. . . . . . . . . Astegopteryx bambusae/minuta/singaporensis/unimaculata;
see key to Bambusa aphids
SIPH pores on small, pigmented cones without hairs. Aptera with wax
pore plates (when present) arranged in groups . . . . . . . . . . . . . . . . . . . 5
5 Wax pore plates restricted to ABD TERG 6, 7 and 8 (except occasionally
one on marginal sclerites of ABD TERG 5). Large, rather elongate aphid,
BL usually more than 2.5 mm . . . . . . . . . . . . . Pseudoregma alexanderi
Small groups of spinal and marginal wax pore plates often present on
head, thorax and all abdominal segments. Body oval, less than 2.5 mm
. . . . . . . . . . . . . . . . Pseudoregma albostriata/bambusicola/dendrocalami

DIMORPHANDRA Leguminosae

Dimorphandra mollis Aphis gossypii

DIOSPYROS Ebonies Ebenaceae

[Aphis diospyri Thomas, 1879]

166 Dipterocarpus

Aulacorthum (Neomyzus)
circumflexum
Myzus persicae
(Use key to polyphagous aphids, p. 532)

DIPTEROCARPUS Dipterecarpaceae

Dipterocarpus tuberculatus Cervaphis schouteniae

Dipterocarpus sp. Greenidea sutepensis
(Dipterocarpaceae appear almost immune to attack by aphids.)

Key to distinguish the two species recorded from Dipterocarpus

SIPH with many long stout hairs. Body hairs not on branched processes
............................................ Greenidea sutepensis
SIPH with a subapical ring of small hairs and a few other hairs more
basad. Body with long branched hair-bearing processes (as in Fig. 92A)
........................................... Cervaphis schouteniae

DISTYLIUM Hamamelidaceae

Distylium spp. are primary hosts for many members of the Nipponaphidini,
a tribe in the subfamily Hormaphinae. Takahashi (1962a) and Sorin (1987b)
reviewed the species of this group that form galls on Distylium in Japan; Sorin
illustrated the galls of five species, and the embryos from emigrant alatae
of 12 species, but there are probably numerous other species in East and
Southeast Asia still undescribed. Determinations using the partial key below
should therefore be treated very cautiously.

Host Plant List

Distylium racemosum Dinipponaphis autumna
(Isunaki) Metanipponaphis cuspidatae,
[rotunda]
Monzenia globuli
Neothoracaphis hangzhouensis,
yanonis
Nipponaphis distychii, distyliicola,
[litseae], [machili(?)], monzeni
Nipponaphis sp. (Sorin, 1987b)
 Distylium 167

Quadratus yoshinomiyai
Sinonipponaphis monzeni
D. stellare Distylaphis foliorum
Greenidea flacourtiae
Neohormaphis calva
Reticulaphis distylii
Schizoneuraphis gallarum,
longisetosa
Toxoptera aurantii

Key to the aphids on Distylium

1 Apterae in free-living colonies on the tree . . . . . . . . . . . . . . . . . . . . . . . 2
- Alatae emerging from galls (other morphs not keyed) . . . . . . . . . . . . . 4
2 SIPH pore-like. Antennae very short, 3- or 4-segmented. Wax pore plates
well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Distylaphis foliorum
- SIPH tubular. Antennae 6-segmented. Wax pore plates not developed
............................................................. 3
3 SIPH broadest at midlength, with numerous long hairs. CAUDA short
and broad-based, with an apical papilla. No stridulatory apparatus
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea flacourtiae
SIPH tapering from base to flange, without hairs. CAUDA tongue-
shaped, rounded at apex. Stridulatory apparatus present (Fig. 122) ...
.............................................. Toxoptera aurantii
4 Antennae 4-segmented (Fig. 45A). Abdominal spiracles present on seg-
ments 2-6 (i.e. 5 on each side of abdomen) . . . . . . . . . . . . . . . . . . . . . . .
........................................ Quadrartus yoshinomiyai
Antennae 5-segmented. Abdominal spiracles only present on segments 2-5
(4 on each side of abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Media of forewing unbranched, united basally with Cula and then again
with Cu lb (Fig. 46A) . . . . . . . . . . . . . . . . . . . . . . . . . Neohormaphis calva
Media of forewing usually once-branched, often basally indistinct or
absent, not united with Cula and Culb . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Basal part of ANT III without rhinaria; distance between most proximal
rhinarium and base of segment more than 1.5 times x longer than ANT
II (Fig. 45B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dinipponaphis autumna
- Distance between most proximal rhinarium on ANT III and base of seg-
ment equal to or less than length of ANT II . . . . . . . . . . . . . . . . . . . . 7
7 Vein Culb in forewing not noticeably thicker than vein Cula (Fig. 46B)
............................................... Monzenia globuli
168 Distylium

Fig. 45. A, antenna of Quadratus yushinomiyai (alata from gall); B, same for Dinipponaphis autumna;
C, same for Nipponaphis distychii; D, abdominal spiracle of Schizoneuraphis longisetosa; E, abdominal
spiracle of Nipponaphis monzeni; F, SIPH of N. monzeni; G, ANT I and II of Distylaphis foliorum alata
from gall; H, same for Nipponaphis distyliicola; I, same for Neothoracaphis yanonis; J, same for
Reticulaphis distylii.

- Vein Cu1b in forewing clearly thicker than vein Cula, especially at base
............................................................. 8
8 Wing membrane heavily infuscated, at least between vein Culb and base
of forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Wing membrane between veins almost hyaline, or slightly but evenly
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
9 Antenna more than 2 x as long as head width across (and including) eyes,
ANT III being as long as or a little longer than fore tibia (Fig. 45C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis distychii
- Antenna less than 2 x as long as head width across eyes. ANT III clearly
shorter than fore tibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
 Distylium 169

Fig. 46. Forewings of alate gallicolae of A, Neohormaphis calva (after Noordam, 1991), B, Monzenia
globuli, C, Distylaphis foliorum, D, Schizoneuraphis longisetosa, E, Nipponaphis distylicola,
F, Schizoneuraphis gallarum, G, Neothoracaphis yanonis, H, Reticulaphis distylii.

10 ABD TERG 8 with 4-8 hairs . . . . . . . . . . Metanipponaphis cuspidatae

- ABD TERG 8 with 10 or more hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Greatest width of ANT II about the same as its greatest length (Fig. 45G).
Tibial hairs maximally about 40 µm, not exceeding width of tibia at mid-
point. Forewing vein Cu1b with a distinct shallow S-curve (Fig. 46C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Distylaphis foliorum
- Greatest width of ANT II at least 1.1 times more than its greatest length
(Fig. 45H). Tibial hairs long and fine, mainly longer than 50 µm and
170 Dodonaea

exceeding width of hind tibia at midpoint. Forewing vein Culb straight or

only slightly curved distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Forewing with a clearly defined hyaline patch at the base of the media,
and vein Cu lb slightly curved distally (Fig. 46D). Abdominal spiracles
with opercular sclerites almost completely surrounding spiracular pores
(Fig. 45D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneuraphis longisetosa
- Forewing membrane often slightly paler at base of media, but no clearly
defined hyaline patch, and vein Culb virtually straight (e.g. Fig. 46E).
Abdominal spiracles with opercular sclerites extending not more than half-
way around spiracular pores (e.g. Fig. 45E) . . . . . . . . . . . . . . . . . . . . . 13
13 BL more than 2.8 mm. SIPH with a black sclerotic section (Fig. 45F), and
with basal diameter about equal to middle diameter of ANT III (including
rhinaria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis monzeni
- BL less than 2.8mm. SIPH pale and inconspicuous, with basal diameter
much less than middle diameter of ANT III (including rhinaria) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis distyliicola
14 Forewing rather narrow, more than 2.5 times longer than its greatest
width, with veins (except media) dark and pterostigma blackish (Fig. 46F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneuraphis gallarum
- Forewing broad, less than 2.5 times longer than its greatest width, with
veins pale (except sometimes Cu lb ) and pterostigma dusky . . . . . . . . 15
15 Forewing veins Cula and Cu lb joined at their bases (Fig. 46G). Greatest
width of ANT II up to about 1.25 times its greatest length (Fig. 45I).
Antennae with annular rhinaria rather widely spaced . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neothoracaphis yanonis/hanzhouensis
- Forewing veins Cu1a and Cu1b separate at their bases (Fig.46H). Greatest
width of ANT II at least 1.35 times its greatest length (Fig. 45 J). Antennae
with annular rhinaria closely spaced . . . . . . . . . . . . Reticulaphis distylii

DODONAEA Sapindaceae

Dodonaea boroniaefoliae Myzus persicae

D. viscosa Aphis gossypii, spiraecola
(Use key to polyphagous aphids, p. 532)
 Duabanga 171

DOMBEYA Sterculiaceae

Dombeya rotundifolia Aphis gossypii

Dombeya sp(p). Myzus persicae
Sitobion africanum

Key to aphids on Dombeya

SIPH of aptera long and black with a subapical zone of polygonal
reticulation. CAUDA long and pale . . . . . . . . . . . . Sitobion africanum
Aptera without this combination of characters . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

DOVYALIS Flacourtiaceae

Dovyalis caffra Aphis spiraecola

Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

DUABANGA Lythraceae

Duabanga sonneratioides Eutrichosiphum sikkimense

Tinocallis himalayensis
Tinocallis (Quednaucallis) distincta

Key to aphids on Duabanga

1 SIPH long, tubular and bearing numerous long hairs in both apterae and
alatae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum sikkimense
SIPH short, truncate cones without hairs. All viviparae alate . . . . . . 2
2 ANT III-VI mainly dark, with III bearing 19-28 secondary rhinaria over
c. 0.8 of length . . . . . . . . . . . . . . . . . Tinocallis (Quednaucallis) distincta
ANT III-VI mainly pale, only ever dark apically; III with 3-17 secondary
rhinaria over 0.4-0.7 of length . . . . . . . . . . . . . . Tinocallis himalayensis
172 Dysoxylum

DYSOXYLUM Meliaceae

Dysoxylum sp. Toxoptera odinae

EHRETIA Boraginaceae

Ehretia acuminata Rhopalosiphoninus ehretis

E. buxifolia Toxoptera aurantii
E. microphylla = E. buxifolia

Key to aphids on Ehretia

SIPH strongly clavate. Antennal tubercles well developed. No stridulatory
apparatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphoninus ehretis
SIPH cylindrical or tapering. Antennal tubercles weakly developed.
Stridulatory apparatus present (Fig. 122) . . . . . . . . . Toxoptera aurantii

ELAEAGNUS Elaeagnaceae

Elaeagnaceae are the primary hosts of the eight to nine species of Capito-
phorus known to have host alternation. One species (shepherdiae) seems to
have a specific primary host association with Shepherdia argentea (which is
not Elaeagnus argentea), but the other species are apparently able to utilize
either Elaeagnus or Hippophae, presumably depending on host availability.

Host Plant List

Elaeagnus angustifolia Capitophorus archangelskii,
(Oleaster, Russian Olive) elaeagni, hippophaes,
[shepherdiae]
[Cryptomyzus korschelti]
E. argentea Capitophorus elaeagni, pakansus
E. canadensis Capitophorus elaeagni
E. commutata Capitophorus hippophaes,
pakansus
 Elaeagnus 173

E. crispa Capitophorus elaeagni, hippophaes

ssp. javanicus
E. glabra Capitophorus elaeagni
E. multiflora Capitophorus elaeagni, hippophaes
(Cherry Elaeagnus) ssp. javanicus
[Myzus gumi Shinji, 1922b]
E. occidentalis Capitophorus archangelskii,
hippophaes
E. oldhami Capitophorus elaeagni
Sinolachnus niitakayamensis
E. pungens Capitophorus elaeagni, hippophaes
ssp. javanicus
E. umbellata Capitophorus elaeagni, hippophaes
ssp. javanicus
[Myzus gumi Shinji, 1922b]
Elaeagnus spp. Capitophorus ?inulae,
meghalayensis, similis

Key to aphids on Elaeagnus and Hippophae (apterae viviparae unless other-

wise stated)
l ANT PT/BASE less than 1. SIPH on broad dark hairy cones. Dorsal hairs
with pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 1. SIPH long and tubular. Dorsal hairs with
expanded apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 R IV tapering, only about half as wide at its junction with R V as at its
base; R V longer than its basal width. Alata with very numerous, small
protruberant secondary rhinaria scattered over ANT III and IV . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinolachnus niitakayanensis
- R IV almost cylindrical, not much narrower distally than at base; R V
shorter than its basal width. Alata with c. 10 rather large secondary
rhinaria on ANT III, and 2-3 on IV . . . . . . . . . . . Lachnus wichmanni
3 SIPH distinctly swollen on inner side subapically . . . . . . . . . . . . . . . . . 4
- SIPH cylindrical or tapering, or only very slightly swollen (to less than
1.1 of minimum width) near apex, or (in alata) thinnest in middle and
gradually thicker towards base and apex . . . . . . . . . . . . . . . . . . . . . . . . 6
4 ABD TERG 1-4 usually without submarginal hairs, so that there are only
3 complete longitudinal rows of hairs on each side; these hairs have fan-
shaped apices and are usually longer than their maximum width. Marginal
hairs usually single. SIPH less than 2.7 x CAUDA. Dorsal cuticle not
variolate or reticulate . . . . . . . . . . . . . . . . . . . . Capitophorus hippophaes
174 Elaeagnus

- ABD TERG 1-4 usually with submarginal hairs, so that there are 4
more-or-less complete longitudinal rows of hairs on each side; these hairs
have mushroom- or balloon-shaped apices, and are shorter than their
maximum widths. Marginal hairs usually duplicated. SIPH more than
2.7 x CAUDA. Dorsal cuticle with variolate or reticulate sculpturing
............................................................. 5
5 ABD TERG 1-4 with pleural hairs usually single. BL probably more
than 1.4mm, and ANT PT/BASE probably more than 2.5 except in
fundatrices. (The spring generations of this species on its primary host
are not yet recorded) . . . . . . . . . . . . . . . . . . . . . . . . Capitophorus xanthii
- ABD TERG 1-4 with pleural hairs mostly duplicate. BL less than 1.4 mm,
ANT PT/BASE less than 2.0 . . . . . . . . . . . . Capitophorus shepherdiae
6 ABD TERG 1-4 each with 6-8 hairs (usually only one pair each of spinal,
pleural and marginal hairs) . . . . . . . . . . . . . . . . . . Capitophorus elaeagni
- ABD TERG 1-4 each with 10-26 hairs, due to duplication or multiplica-
tion of spinal, pleural and marginal hairs on each tergite . . . . . . . . . 7
7 SIPH short, tapering continuously from base to apex, less than 11 x
longer than width at midpoint . . . . . . . . . . . Capitophorus himalayensis
- SIPH attenuated, almost cylindrical except at base, or with slight sub-
apical swelling; more than 12 x width at midpoint . . . . . . . . . . . . . . . 8
8 Hairs on ANT III all short, thin and inconspicuous, 0.3-0.5 x basal
diameter of segment. R IV+V 2.5-3.1 x HT II . . . . . . . . . . . . . . . . . . . .
.................................... Capitophorus meghalayensis*
- ANT III with 1-5 forwardly-directed thick clavate hairs like those on ANT
II or a little shorter, 0.5-1.2 x longer than base of segment. R IV+V
1.5-2.6 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Thick capitate hairs on ANT III all less than 0.8 x basal diameter of
segment. (R IV+V 1.5-2.1 x HT II) . . . . . . . . . . . Capitophorus similis
- Longest capitate hairs on ANT III more than 0.8 x basal diameter of
segment. (R IV+V 1.6-2.6 x HT II) . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 ABD TERG 1-4 each with 18-26 capitate hairs. R IV+V 2.1-2.6 x HT
II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capitophorus pakansus
- ABD TERG 1-4 each with 12-19 capitate hairs. R IV+V 1.6-2.1 x HT
II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Capitophorus archangelskii
 Eleutherococcus 175

ELAEIS Palmae

Elaeis guineensis Astegopteryx nipae

(For palm aphids see B & E, 1984)

ELAEOCARPUS Elaeocarpaceae

Elaeocarpus japonicus Eutrichosiphum pasaniae

E. serratus Acyrthosiphon elaeocarpi
E. sikkimensis [Trichaitophorus recurvispinosus]

Key to aphids on Elaeocarpus

- SIPH with numerous long hairs. R IV+V long and narrow . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pasaniae
- SIPH without hairs. R IV+V short . . . . . . . . Acyrthosiphon elaeocarpi

ELEUTHEROCOCCUS Araliaceae

Eleutherococcus spp. Aphis acanthopanaci, gossypii,

spiraecola

Key to aphids on Eleutherococcus

1 CAUDA paler than SIPH. Hind femur with all hairs clearly shorter than
its basal diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
CAUDA as dark as SIPH. Hind femur with long, fine hairs, at least some
of which are longer than its basal diameter . . . . . . . . . . . . . . . . . . . . . . 2
2 Lateral tubercles present on most of abdominal segments 2-4, as well as
on 1 and 7. R IV+V more than 1.25 times longer than HT II .......
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis acanthopanaci
No lateral tubercles on abdominal segments 2-4. R IV+V less than 1.2
times longer than HT II . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
176 Encephalartos

ENCEPHALARTOS Zamiaceae

Aphis gossypii

ENGELHARDTIA Juglandaceae

Engelhardtia spicata Aiceona himalaica

[Greenidea (Trichosiphum) heeri]
Kurisakia indica

Key to aphids on Engelhardtia

Antennae 5-segmented, less than 0.5 of BL. SIPH cones bearing less than
10 long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kurisakia indica
Antennae 6-segmented, more than 0.5 of BL. SIPH cones bearing
numerous short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona himalaica

ERIOBOTRYA Rosaceae

Host Plant List

Eriobotrya dubia Nippolachnus bengalensis
Tuberolachnus sclerata
E. japonica Aphis eugeniae, gossypii, pomi,
(Japanese Medlar, Loquat) spiraecola
Brachycaudus persicae
Hyalomyzus eriobotryae
Nippolachnus piri,
[xitianmushanus]
Pyrolachnus [macroconus], pyri
Tuberolachnus sp. nr sclerata
E. petiolata Nippolachnus himalayensis
Prociphilus sp. near himalayaensis
Tuberolachnus sclerata
 Ervatamia 177

Key to aphids on Eriobotrya (a revised and extended version of the key in

B & E, 1984)
1 ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2 Body densely hairy. SIPH as large pores on hairy cones . . . . . . . . . . 3
Body sparsely hairy. SIPH not on hairy cones . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus sp. near himalayaensis
3 Legs pale. Body elongate oval, at least 2 x its maximum width .... 4
Legs mainly dark. Body broadly oval, less than 2 x its maximum width
............................................................. 5
4 ANT PT/BASE 0.25-0.35, without any fine hairs distal to primary
rhinarium. R IV+V 0.14-0.18 mm long, 0.62-0.75 X HT II, and bearing
9-16 accessory hairs . . . . . . . . . . . . . . . . . . . . . Nippolachnus bengalensis
- ANT PT/BASE 0.45-0.75, with some long fine hairs distal to primary
rhinarium. R IV+V 0.17-0.22 mm long, 0.7-1.0 x HT II, and bearing
more than 20 accessory hairs . . . . . . . . . . . . . . . . . . . . Nippolachnus piri
5 Eye without an ocular tubercle. Hairs on antennae and frons very long
and fine, maximally 3 or more x basal diameter of ANT III . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nippolachnus himalayensis
Eye with a prominent ocular tubercle. Hairs on antennae and frons max-
imally 2.5 x basal diameter of ANT III or shorter . . . . . . . . . . . . . . . 6
6 ABD TERG 4 with a large spinal tubercle . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Tuberolachnus (Tuberolachniella) sclerata
ABD TERG 4 without a large spinal tubercle . . . . . . Pyrolachnus pyri
7 Dorsal abdomen black. CAUDA shorter than its basal width in dorsal
view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus persicae
Dorsal abdomen pale. CAUDA longer than its basal width . . . . . . . . 8
8 Antennal tubercles well developed, with inner faces convergent in dorsal
view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalomyzus eriobotryae
Antennal tubercles poorly developed . . . . . . . . . . . . . . . . . . . Aphis spp.
(use couplets 21-25 in key to aphids on Malus)

ERVATAMIA = TABERNAEMONTANA
178 Erythrina

ERYTHRINA Leguminosae

Erythrina ovalifolia Toxoptera citricidus

Erythrina spp. Myzus persicae
(Use key to polyphagous aphids on p. 532)

ESCALLONIA Escalloniaceae

Escallonia pulverulenta Myzus persicae

EUCALYPTUS Myrtaceae

Host Plant List

Eucalyptus camaldulensis Aphis fabae, gossypii
Myzus persicae
Toxoptera aurantii
Eu. gomphocephala Toxoptera aurantii
Eu. grandis Aphis fabae, gossypii
Eu. macarthurii Macrosiphum euphorbiae
Eu. melliodora Aphis gossypii
Eu. robusta Toxoptera aurantii, odinae
(Use key to polyphagous aphids, p. 532)

EUCLEA Ebenaceae

Euclea crispa Aphis gossypii

 Eugenia 179

EUCOMMIA Eucommiacee

Eucommia ulmoides Myzus persicae

EUGENIA Myrtaceae

A small but diverse aphid fauna occurs on Eugenia, including polyphagous

species and others specific to Myrtaceae. Most records do not identify the host
to species.

Host Plant List

Eugenia aquea Aphis gossypii
Toxoptera aurantii
Eu. densiflora Aphis gossypii
Sinomegoura citricola
Eu. jambolana Greenidea (Trichosiphum) heeri
Eu. mooneaca (mooniana?) Toxoptera aurantii
Eugenia spp. Aphis eugeniae
Aulacorthum solani
Chaitophorus pakistanicus
Greenidea (Trichosiphum)
formosana
Neotuberaphis indica
Taiwanaphis decaspermi

Key to aphids on Eugenia

1 ANT PT/BASE less than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Aptera with a single pair of tubercles on the head, each bearing a
spine-like hair, 3-faceted eyes, and with a row of marginal wax glands
around the body. Alata with numerous annular rhinaria on ANT III, IV
and V, and forewing veins not heavily bordered . . . . . . . . . . . . . . . . . . .
............................................ Neotuberaphis indica
Aptera without tubercles on head bearing spine-like hairs, with multi-
faceted eyes, and body without a row of marginal wax glands. Alata with
oval or transversely elongate secondary rhinaria confined to ANT III, and
180 Euonymus

forewing veins heavily bordered with fuscous . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis decaspermi
3 Dorsal body hairs of aptera all very long, often with bifurcate or multi-
furcate apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Dorsal body hairs all short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
4 SIPH long, dark, tubular and covered with long hairs like those on
dorsum. R IV+V about 2 x HT II . . . . . . Greenidea formosana/heeri
- SIPH short, pale, truncate and without hairs. R IV+V about 0.8 x HT
II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus pakistanicus

EUONYMUS Celastraceae

Several subspecies or sibling species of the Aphis fabae group use Euonymus
europaeus as their primary host in Europe (Müller and Steiner, 1986). These
are suspected to hybridize in nature to some extent, and the morphological
separations attempted in couplets 6 and 7 of the key below may not be very
reliable. For biological differences between members of this group consult the
text (p. 559).

Host Plant List

Euonymus alata Aphis fabae group
Aulacorthum magnoliae
Eu. alata var. ciliatodentatus [Myzus komaumii Shinji, 1943]
Eu. atropurpurea Aphis fabae group
Eu. europaeus Aphis fabae group
(cirsiiacanthoidis, euonymi,
fabae, solanella)
Macrosiphum euphorbiae
Eu. japonicus Aphis fabae group
Aulacorthum magnoliae
Myzus persicae
Eu. sieboldianus Aulacorthum magnoliae
Euonymus sp(p). (unidentified) Toxoptera aurantii

Key to aphids on Euonymus

1 Antennal tubercles well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Antennal tubercles weakly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
 Euonymus 181

2 Antennal tubercles with inner faces convergent in dorsal view (Fig. 42I).
Aptera without any secondary rhinaria on ANT III. Alata with a dark
dorsal abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
Antennal tubercles with inner faces parallel or divergent in dorsal view.
Aptera with one or more secondary rhinaria near base of ANT III. Alata
either without dorsal abdominal markings or with transverse segmentally
divided bars ................................................. 3
3 Antennal tubercles smooth with inner faces divergent in dorsal view
(Fig. 42K). SIPH tapering, with a longer-than-wide subapical zone of
polygonal reticulation. (Longest hairs on ANT III more than 0.5 of basal
diameter of segment.) Alata without dorsal abdominal markings .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
Antennal tubercles with inner faces scabrous, parallel in dorsal view
(Fig. 42J), SIPH without a zone of polygonal reticulation. Alata with
brown transverse bars on dorsal abdomen . . . . . . . . . . . . . . . . . . . . . . . 4
4 Femora and CAUDA mainly dark. SIPH swollen on distal half . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
Femora and CAUDA mainly pale. SIPH tapering . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
5 ANT PT/BASE 3.5-5.0. Abdominal sternites 5 and 6 with conspicuous
sclerotic ridges, and hind tibia with a row of evenly spaced, short,
stridulatory pegs (Fig. 122) . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
ANT PT/BASE 1.7-3.5. Stridulatory ridges and pegs absent . . . . . . 6
6 ANT III 3-5 x longer than the longest hair borne upon it, which is
2-3 x basal diameter of the segment. Marginal tubercles completely
absent from ABD TERG 2-6. Males apterous, oviparae with hind tibiae
only slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis euonymi
ANT III 4-20 x longer than the longest hair borne upon it, which is
0.6-2.5 (but rarely more than 2.0) X basal diameter of the segment. ABD
TERG 2-6 with a total of 0-7 (usually 1-6) marginal tubercles. Males
alate, oviparae with strongly swollen hind tibiae . . . . . . . . . . . . . . . . . . 7
7 ANT III of aptera 11-20 x longer than the longest hair borne upon it.
Hairs on marginal sclerites of ABD TERG 3 of aptera 12-49 µm long, of
alata 18-51µm long, usually less than 2 x maximum diameter of the
spiracular pore on that segment . . . . . . . . . . . . . . Aphis fabae solanella
ANT III of aptera 4-9 x longer than the longest hair borne upon it. Hairs
on marginal sclerites of ABD TERG 3 of aptera 40-85 µm long, of alata
31-83 µm long, more than 2 x maximum diameter of spiracular pore on
that segment . . . . . . . . . . . . . Aphis fabae fabae/fabae cirsiiacanthoidis
182 Euodia

EUODIA = EVODIA Scop. nec. Gaertn. Rutaceae

Euodia triphylla Acyrthosiphon evodiae

Toxoptera aurantii
Key to aphids on Euodia
Antennal tubercles well developed. No stridulatory apparatus . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon evodiae
Antennal tubercles weakly developed. Stridulatory apparatus present
(Fig. 122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
(or try key to polyphagous aphids, p. 532)

EUPHORBIA (trees only) Euphorbiaceae

Euphorbia candelabrum Myzus persicae

Euphorbia spp. (African) Sitobion phyllanthi

Key to aphids on Euphorbia

SIPH pale or dusky, slightly clavate. Antennal tubercles well developed,
with inner faces convergent in dorsal view (Fig. 42I) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
SIPH dusky or dark, tapering or cylindrical. Antennal tubercles weakly
developed, with inner faces broadly divergent in dorsal view . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion phyllanthi
(or try key to polyphagous aphids, p. 532)

EUPTELEA Eupteleaceae

Euptelea polyandra Myzus persicae

EUSCAPHIS Staphyleaceae

Euscaphis japonica Aphis horii

 Fagus 183

FAGARA = ZANTHOXYLUM

FAGUS Beeches Fagaceae

Host Plant List

Fagus crenata Mesocallis fagicola
Phyllaphis fagifoliae
Platyaphis fagi
F. grandifolia Eriosoma crataegi group
Grylloprociphilus imbricator
Longistigma caryae
Phyllaphis grandifoliae
F. japonica Phyllaphis fagifoliae
F. sylvatica Lachnus pallipes
(European Beech) Phyllaphis fagi
Fagus sp. Stomaphis fagi

Key to aphids on Fagus

1 Aptera with margins of body flattened and coarsely serrate like the teeth
of a circular saw, each 'tooth' with a backwardly-directed point bearing
a minute hair (Fig. 47A). Neither rostrum nor antennae exceeding 0.2 mm
in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platyaphis fagi
If apterae occur then they are without flattened, serrate margin to body,
and the rostrum and antennae both exceed 0.2 mm in length . . . . . . 2
2 Antennae densely hairy, with length of hairs mostly equal to or exceeding
basal diameter of ANT III. SIPH on broad dark hairy cones . . . . . . 3
Antennae with sparse, short hairs. SIPH if present then not as broad
dark hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 Rostrum longer than body. HT II nearly twice as long as segment II
of fore- and mid-tarsi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis fagi
Rostrum much shorter than body. HT II only a little longer than segment
II of fore- and mid-tarsi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 ANT VI BASE about 3 x longer than wide. Forewings of alata uniformly
fuscous, with pterostigma extending around tip of wing. BL more than
4.7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
184 Fagus

Fig. 47. A, margin of abdomen of Platyaphis fagi (aptera); B, dorsal abdominal wax pore plate of
fundatrix of Grylloprociphilus imbricator; C, dorsal abdominal wax pore plate of Eriosoma crataegi,
D, ANT VI of Phyllaphis fagi (aptera); E, dorsal abdominal wax pore plate of Ph. fagi (alata).

ANT VI short and stubby, its base less than 2 x longer than wide.
Forewings of alata maculate, with pterostigma not extending to tip of
wing. BL usually less than 4.5 mm . . . . . . . . . . . . . . . . Lachnus pallipes
5 Apterae with faceted wax glands. Alatae with secondary rhinaria on ANT
III-V or III-VI, media unbranched or once-branched, and a broadly
rounded CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Apterae (if produced) with wax glands comprising groups of ring-shaped
pores (Fig. 47E). Alatae with secondary rhinaria only on ANT III, a
twice-branched media and a knobbed CAUDA . . . . . . . . . . . . . . . . . . . 7
6 Apterae (fundatrices) with 4-segmented antennae and circular groups of
wax pore plates on all segments, each with large central facets surrounded
by very much smaller marginal ones (Fig. 47B). SIPH absent. Alata with
strongly ciliated secondary rhinaria and an unbranched media .......
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grylloprociphilus imbricator
Apterae with 6-segmented antennae, wax pore plates smaller and usually
comprising a ring of large facets around a central area which may be
 Ficus 185

divided into small cells (Fig. 47C). SIPH present as partly sclerotized rings
with surrounding hairs. Alatae with unciliated secondary rhinaria and a
once-branched media . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma crataegi
7 All viviparae alate. ANT PT/BASE about 1. Dorsal abdomen of alata
pale. SIPH as pale, truncate cones . . . . . . . . . . . . . . Mesocallis fagicola
Viviparae apterous or alate. ANT PT/BASE less than 0.25 (Fig. 47D).
Dorsal abdomen of alata pale or with variably developed transverse bars
and pigmented marginal sclerites bearing numerous wax pores (Fig. 47E).
SIPH pore-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Alatae with pale head and pterothorax and no dorsal abdominal cross-
bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaphis fagifoliae
Alatae with dark head and pterothorax and pigmented dorsal abdominal
cross-bands and marginal sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaphis fagi (and grandifoliae?)

FICUS Figs Moraceae

Aphids on Ficus carica are keyed in B & E (1984). The present treatment is
extended to cover all aphids recorded from the genus Ficus.

Host Plant List

Ficus akeotsang Toxoptera aurantii
F. ampelos Greenidea ficicola
Toxoptera aurantii
F. aurantiaca Toxoptera aurantii
F. bengalensis Aphis gossypii, spiraecola
(Banyan Tree, Bargat) Greenidea ficicola
Reticulaphis distylii group
F. benjamina Greenidea artocarpi
Nipponaphis ficicola
Reticulaphis distylii group
F. carica Aphis fabae, kachkouli, gossypii
(Common Fig) Greenidea ficicola
Greenidea (Trichosiphuni)
guangzhouensis
Reticulaphis distylii fid
Sitobion africanum
Toxoptera aurantii
186 Ficus

F. chlorocarpa Greenidea ficicola

F. dekdekena Sitobion africanum
F. depressa Reticulaphis distylii group
F. diupacea Reticulaphis distylii group
F. elastica Aphis gossypii, spiraecola
Reticulaphis distylii group
Toxoptera citricidus
F. foveolata Reticulaphis distylii foveolata
F. glomerata Greenidea artocarpi, ficicola
F. hispida Toxoptera aurantii
F. ingens Aphis gossypii
Sitobion africanum
Toxoptera aurantii, citricidus
F. iteophylla Myzus persicae
F. johannis Aphis kachkouli
F. macrophylla Greenidea ficicola
F. microcarpa Aphis gossypii, spiraecola
Greenidea ficicola
Reticulaphis distylii
Toxoptera schlingen
F. mucuso Aphis craccivora
F. nitida var. hillyi Greenidea ficicola
F. obscura Sinomegoura citricola
Toxoptera aurantii
F. pseudopalma Reticulaphis distylii group
F. pumila Aphis spiraecola
Aulacorthum (Neomyzus)
circumflexum
F. religiosa Aphis craccivora, gossypii,
(Bo-Tree, Peepul) spiraecola
Toxoptera aurantii
F. retusa Greenidea ficicola
Greenidea (Trichosiphum) fici,
formosana
Reticulaphis distylii
F. ribes var. cuneata Matsumuraja calorai
 Ficus 187

F. septica Reticulaphis distylii

F. sycomorus Aphis gossypii, spiraecola
F. thonningii Sitobion africanum
F. vallis Sitobion halli
F. variegata Greenidea ficicola
F. wightiana Greenidea (Trichosiphum)
formosana
Reticulaphis distylii group
Toxoptera odinae
Ficus sp. Lachnus fici

Key to aphids on Ficus

1 Aptera aleyrodiform or coccidiform, with sclerotic dorsum, eyes of only
3 facets and short, at most 3-segmented, antennae. Alata with annular
rhinaria on the antenna and an unbranched or once-branched media
in the forewing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Aptera of the normal aphid form, eyes multifaceted and antennae 5- or
6-segmented. Alata with circular or oval antennal rhinaria and forewing
with once- or twice-branched media . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Aptera with SIPH, and with dorsal cuticle densely covered with trans-
parent 'pustules' (Fig. 48A). Marginal hairs with acute apices . . . . . . . .
............................................ Nipponaphis ficicola
- Aptera without SIPH, and with dorsal cuticle very dark, heavily sclerotic
and reticulated (Fig. 48C). Marginal hairs with acute, blunt or fan-shaped
apices ....................................................... 3
3 Aptera with a pair of long, stout, flattened, apically rounded hairs on the
median area between the eyes (Fig. 48B), like those along margins of
body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reticulaphis mirabilis
- Aptera with all mediodorsal hairs, including those between eyes, minute;
much shorter than marginal and frontal hairs, which have either acute or
flattened (e.g. Fig. 48C) apices . . . . . . . . . . . Reticulaphis distylii group
4 ANT PT/BASE less than 0.5. SIPH as pores on dark hairy cones ...
.................................................. Lachnus fici*
- ANT PT/BASE more than 1. SIPH tubular . . . . . . . . . . . . . . . . . . . . . 5
5 SIPH covered with long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
6 SIPH of aptera reticulated only at the base, and rather densely spinulose
over most of length (Fig. 48D). (Alata with 18-26 secondary rhinaria on
ANT III) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea formosana group
 188 Ficus

Fig. 48. A, SIPH and adjacent cuticle of Nipponaphis ficicola (aptera); B, head of Reticulaphis mirabilis
(after Takahashi, 1939a); C, head of Reticulaphis distylii ssp. minutissima (with detail of dorsal cuticle);
D, SIPH of Greenidea formosana group (aptera); E, SIPH of G. ficicola (aptera); F, dorsal view of front of
head and processes on ANT I of Matsumuraja calorai aptera; 6, ANT I-IV of Toxoptera schlingen alata;
H, ANT I-IV of T. aurantii alata.

- SIPH of aptera reticulated over most of length, spinulose only on distal

part (Fig.48E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 SIPH of aptera longer than hind tibiae and a little more than 0.5 x BL.
Alata with 28-35 secondary rhinaria on ANT III . . . . . . . . . . . . . . . . . . .
............................................. Greenidea artocarpi
 Ficus 189

- SIPH of aptera usually a little shorter than hind tibiae and clearly less than
0.5 x BL. Alata with 12-22 secondary rhinaria on ANT III . . . . . . . . .
.............................................. Greenidea ficicola
8 ANT I of aptera very broad, with a finger-like distal projection, extending
beyond apex of ANT II and bearing short, capitate hairs (Fig. 48F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Matsumuraja calorai
- ANT I without a finger-like projection . . . . . . . . . . . . . . . . . . . . . . . . 9
9 SIPH with a subapical zone of polygonal reticulation . . . . . . . . . . . . 10
- SIPH without any polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . 11
10 Aptera with SIPH usually uniformly dark, and dorsal abdomen often with
a pattern of dark segmental markings. R IV+V 0.8-0.9 x HT II. Alata
with 4-14 secondary rhinaria mostly concentrated on basal half of ANT
III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion africanum
- Aptera with SIPH often pale at base, and dorsal abdomen without
dark markings. R IV+V 0.95-1.15 X HT II. Alata with 14-18 second-
ary rhinaria in spaced-out row along most of ANT III . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion halli
11 Antennae always 5-segmented, in both apterae and alatae . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis kachkouli
- Antennae 6-segmented (except sometimes in small apterae of Aphis
gossypii) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Stridulatory apparatus present, comprising stridulatory ridges on abdom-
inal sternites 5 and 6 and evenly-spaced peg-like hairs on the hind tibiae
(Fig. 122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Stridulatory apparatus absent ... go to key to polyphagous aphids, p. 532
13 SIPH 0.4-0.7 x CAUDA, and 0.05-0.08 x BL . . . . Toxoptera odinae
- SIPH 0.9-1.9 x CAUDA, 0.11-0.18 X BL . . . . . . . . . . . . . . . . . . . . . . 14
14 CAUDA with 19-54 hairs (rarely less than 25). Hairs on ANT III of aptera
mostly longer than basal diameter of segment. Alata with forewing media
usually twice-branched . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera citricidus
- CAUDA with 7-26 hairs (rarely more than 20). Hairs on ANT III of
aptera mostly shorter than basal diameter of segment. Alata with forewing
media usually once-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Alata with 12-18 large and small secondary rhinaria on ANT III, 3-6 on
ANT IV and 0-2 on ANT V (Fig. 48G). ANT PT/BASE often less than
3.5 in aptera, and often less than 3.75 in alata . . . . . . . . . . . . . . . . . . . .
............................................ Toxoptera schlingeri
- Alata with 2-10 more evenly-sized secondary rhinaria on ANT III, 0-3
on ANT IV, 0 on ANT V (Fig. 48H). ANT PT/BASE greater than 3.5
in aptera, greater than 3.75 in alata . . . . . . . . . . . . . Toxoptera aurantii
190 Fraxinus

FRAXINUS Ashes Oleaceae

Fraxinus species are primary or only hosts for several members of the
pemphigine genus Prociphilus, the spring generations of which form leaf
nests. Only the alatae produced by these leaf-nest colonies are keyed here.
Of the other aphids described or recorded from Fraxinus, only one (Myzus
beybienkoi) has been collected more than once. The others are not included
in the key and are indicated by square brackets in the following list.

Host Plant List

Fraxinus americana [Phylloxera (?) fraxini Stebbing,
(White Ash) 1910]
Prociphilus americanus,
fraxinifolii, oriens, pergandei,
probosceus
F. dipetala Prociphilus americanus
F. excelsior [Aphis excelsioris Dahlbom, 1851]
(European Ash) [Aphis gossypii]
Prociphilus americanus, bumeliae,
fraxini, oriens
[Pterocomma fraxini Theobald,
1929]
F. floribunda Prociphilus sp.
F. insularis (?) Prociphilus formosanus
F. latifolia Prociphilus americanus,
fraxinifoliae
F. longicuspis [Chaitophorus fraxinicolus
Matsumura, 1919]
[Siphocoryne fraxinicola
Matsumura, 1917]
F. mandshurica Prociphilus oriens
F. nigra Prociphilus fraxinifolii
F. oregona (= latifolia)
F. oxycarpa [Eucallipterus tiliae]
Prociphilus sp. near fraxini
F. pennsylvanica Prociphilus americanus,
fraxinifolii, probosceus
 Fraxinus 191

F. potamophila Myzus beybienkoi

F. quadrangulata Prociphilus fraxinifolii
F. rhynchophylla Prociphilus oriens
F. sambucifolia Prociphilus fraxinifolii
F. sieboldiana Prociphilus oriens
F. spaethiana Prociphilus oriens
F. syriaca [Aphis gossypii]
F. uhdei Prociphilus fraxinifolii
F. velutina Prociphilus fraxinifolii, oriens
Fraxinus sp. [Prociphilus cheni]

Key to aphids on Fraxinus

1 SIPH tubular, slightly swollen. ANT PT/BASE about 3. All antennal and
dorsal body hairs short with expanded apices. Alata with a black dorsal
abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus beybienkoi
- SIPH absent or only present as small pores. ANT PT/BASE less than
0.5. Antennal and body hairs of variable length, finely pointed. (Alata
with unpigmented abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Apterae in colonies at base of trunk or on roots . . . . . . . . . . . . . . . . . 3
- Alatae emerging from leaf-nest galls in spring or early summer . . . . . . 4
3 BL more than 4 mm. Rostrum distinctly longer than body. R IV+V with
about 50 accessory hairs . . . . . . . . . . . . . . . . . . . Prociphilus probosceus
- BL less than 3 mm. Rostrum very short, R IV+V with 2 accessory hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus fraxinifolii
4 ANT VI BASE with secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . 5
- ANT VI BASE without secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . 7
5 ANT VI BASE with 1-5 irregularly-shaped secondary rhinaria, differing
from those on ANT III (Fig. 49A) . . . . . . . . . . . Prociphilus fraxinifolii
- ANT VI BASE with 8-15 narrow transverse secondary rhinaria like those
on ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 ANT II much longer than broad, with more than 25 hairs; ANT BASE
VI with 10-18 hairs (Fig. 49B). R IV+V with more than 10 accessory hairs.
Head with a posterior dorsal pair of wax pore plates that are larger than
the ocelli (Fig. 49D) . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus probosceus
- ANT II not longer than broad, with less than 10 hairs; ANT BASE VI with
3-6 hairs (Fig. 49C), R IV+V with 4-8 accessory hairs. Head with dorsal wax
pore plates absent or indistinct . . . . . . . . . . . . . . . . Prociphilus pergandei
192 Fraxinus

Fig. 49. A, antenna of Prociphilus fraxinifolii alata; B, antenna of P. probosceus alata; C, ANT I-III and
VI of P. pergandei alata; heads of spring migrant alatae of D, P. probosceus, E, P. bumeliae and
F, P. fraxini (o = ocellus, w = wax gland).

7 ANT V with 4-11 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

- ANT V with 0-2 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 SIPH present as small pores . . . . . . . . . . . . . . . Prociphilus formosanus
- SIPH absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
9 ANT III over 5 x ANT II, as long as or longer than ANT IV+V together.
Head with a pair of large posterior dorsal wax pore plates, conspicuous
as clearly defined pale areas much larger than ocelli (Fig. 49E), but
without anterior dorsal wax pore plates . . . . . . . . Prociphilus bumeliae
- ANT III usually less than 5 x ANT II, usually a little shorter than ANT
IV+V together. Head without or with ill-defined posterior dorsal wax pore
plates, not forming demarcated pale areas. With or without anterior dor-
sal wax pore plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 R IV+V 0.50-0.55 x HT II. Head usually with a pair of small clear
anterior wax pore plates (Fig. 49F). Embryos (inside mother's abdomen)
 Gigantochloa 193

with spinal, pleural and marginal hairs all of similar length . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus fraxini
- R IV+V 0.56-0.72 x HT II. Head usually without any anterior dorsal wax
pore plates. Embryos with only marginal hairs long, about 5 times longer
than the inconspicuous pleural and spinal hairs . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus americanus

FUCHSIA Onagraceae

Aulacorthum solani
Aulacorthum (Neomyzus)
circumflexum
Macrosiphum euphorbiae
Myzus ornatus, persicae
(Use key to polyphagous aphids, p. 532)

GARCINIA Guttiferae

Garcinia multiflora Toxoptera odinae

GARDENIA Rubiaceae

Aphis gossypii, spiraecola

Myzus persicae
Sinomegoura citricola
Toxoptera aurantii, citricidus,
odinae
(Use key to polyphagous aphids, p. 532)

GIGANTOCHLOA Gramineae

Gigantochloa apus Astegopteryx minuta, pallida,

unimaculata
194 Gleditsia

Ceratoglyphina bambusae
Pseudoregma bambusicola
(Use key to aphids on Bambusa, p. 83)

GLEDITSIA Leguminosae

Brachycaudus helichrysi
Myzus persicae
(Use key to polyphagous aphids, p. 532)

GLIRICIDIA Leguminosae

Aphis craccivora

GLOCHIDION Euphorbiaceae

Aphis eugeniae, spiraecola

[Eutrichosiphum makii]
Micromyzus hangzhouensis
Schoutedenia ralumensis

Key to aphids on Glochidion

1 ABD TERG 7 with a pair of large backward-pointed processes. Antenna
5-segmented, ANT PT/BASE less than 1. Aptera with 3-faceted eyes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schoutedenia ralumensis
- ABD TERG 7 without large processes. Antennae 6-segmented, ANT
PT/BASE more than 1. Aptera with multifaceted eyes . . . . . . . . . . . . 2
2 Antennal tubercles well developed. Aptera with 1-3 secondary rhinaria on
ANT III. ANT PT/BASE more than 4 . . . . . . . . . . . . . . . . . . . . . . . . . . .
...................................... Micromyzus hangzhouensis
- Antennal tubercles weakly developed. Aptera without secondary rhinaria
on ANT III. ANT PT/BASE less than 4 . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Hind tibia with some hairs, particularly on basal half, short and peg-like,
clearly differentiated from other tibial hairs. HT I with 3 hairs. Aptera
 Guaiacum 195

with R IV+V 1.15-1.30 x HT II. Alata with SIPH 0.12-0.18 x BL, and
1.4-1.8 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis eugeniae
Hind tibia without any short peg-like hairs. HT I with 2 hairs. Aptera
with R IV+V 1.0-1.16 x HT II. Alata with SIPH 0.09-0.14 x BL and
1.2-1.5 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola

GREVILLEA Proteaceae

Grevillea robusta Aphis gossypii

Myzus persicae
(Use key to polyphagous aphids, on p. 532)

GREWIA Tiliaceae

Grewia occidentalis Toxoptera aurantii

G. oppositifolia Sinomegoura citricola
G. optiva Sinomegoura citricola
G. tiliaefolia Aphis gossypii
Sinomegoura citricola
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

GREYIA Greyiaceae

Greyia sutherlandii Macrosiphum euphorbiae

Toxoptera odinae
(Use key to polyphagous aphids, p. 532)

GUAIACUM Lignum Vitae Zygophyllaceae

No aphids recorded
196 Guazuma

GUAZUMA Sterculiaceae

Guazuma ulmifolia Toxoptera aurantii

GUETTARDA Rubiaceae

Guettarda krugii Aphis spiraecola

HAGENIA Rosaceae

Macrosiphum euphorbiae

HAMAMELIS Hamamelidaceae

Hamamelis japonica Hamamelistes kagamii, miyabei

Hormaphis betulae
H. virginiana Hamamelistes spinosus
Hormaphis hamamelidis

Key to alatae from galls on Hamamelis

1 Antenna 3-segmented. Hind wing with 1 oblique vein (Fig. 50A) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hormaphis hamamelidis
Antenna 5-segmented. Hindwing with 2 oblique veins . . . . . . . . . . . . . 2
2 SIPH inconspicuous or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
SIPH present as pores with sclerotic rims . . . . . . . . . . . . . . . . . . . . . . . 4
3 Hindwing with the 2 oblique veins often incomplete, and well separated
at their bases (Fig. 50B) . . . . . . . . . . . . . . . . . . . . Hamamelistes spinosus
Hindwing with the 2 oblique veins complete; united or close together at
their bases (e.g. Fig. 50C) . . . . . . . . . . . . . . . . . . . . . . Hormaphis betulae
 Harungana 197

Fig. 50. Hindwings of alatae from galls of A, Hormaphis hamamelidis, B, Hamamelistes spinosus,
C, Hormaphis betulae, D, Hamamelistes miyabei.

4 ANT IV distinctly longer than (more than 1.2 x) ANT V. Hindwing with
the 2 oblique veins united or close together at their bases (e.g. Fig. 50D)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamamelistes miyabei
- ANT IV about equal in length to (0.9-1.1 x) ANT V. Hindwing with the
2 oblique veins separated at their bases . . . . . . . Hamamelistes kagamii

HARPEPHYLLUM Anacardiaceae

Harpephyllum caffrum Myzus persicae

Toxoptera aurantii, odinae
(Use key to polyphagous aphids, p. 532)

HARUNGANA Guttiferae

Harungana madagascariensis Aphis gossypii

(Namahasi) Aulacorthum solani
Sitobion congolense, nigeriense
Toxoptera aurantii, odinae

Key to aphids on Harungana

1 SIPH long, with a short subapical zone of polygonal reticulation. CAUDA
long and tapering, that of aptera more than 0.15 x BL. ANT PT/BASE
more than 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
198 Hemiptelea

Fig. 51. Forewing pigmentation of A, Tinocallis coreanus and B, T. takachihoensis.

SIPH long or short, without any polygonal reticulation. CAUDA of

aptera less than 0.13 x BL. ANT PT/BASE less than 6 . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . see key to polyphagous aphids, p. 532
2 SIPH slightly flared apically, with a well-developed flange. Alata without
secondary rhinaria on ANT IV . . . . . . . . . . . . . . . . . Sitobion nigeriense
SIPH narrow or slightly constricted apically, with a very small flange.
Alata sometimes with 1-7 rhinaria on ANT IV . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion congolense

HEMIPTELEA Ulmaceae

Hemiptelea davidii Tinocallis (Pseudochromaphis)

(Simu) coreanus
Tinocallis takachihoensis

Key to aphids on Hemiptelea

Forewing membrane extensively mottled with dark pigment (Fig. 51A).
Head and prothorax pigmented dorsally with a pale spinal stripe and large
 Hibiscus 199

pale areas around the hair bases. Dorsal abdomen without finger-like
tubercles but with dark markings. ANT PT/BASE less than 0.5. ANT III
with about 6-8 widely-spaced oval secondary rhinaria . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis (Pseudochromaphis) coreanus
Forewing membrane with fuscous bordering distal branches of media and
patches at ends of Cula and Cu1b (Fig. 51B). Head and prothorax rather
uniformly dark. Dorsal abdomen pale, with finger-like spinal tubercles on
ABD TERG 1 and 2. ANT PT/BASE about 1 or more. ANT III with
14-18 secondary rhinaria . . . . . . . . . . . . . . . . . Tinocallis takachihoensis

HERNANDIA Hernandiaceae

Hernandia sonora Aphis gossypii

HETEROMELES = PHOTINIA

HIBISCUS Malvaceae

Hibiscus has a very similar aphid fauna to cotton (keyed in Blackman and
Eastop, 1984). The aphids are all oligophagous or polyphagous, so there is
no point in listing the host species separately.

Hibiscus spp. Acyrthosiphon gossypii

[Aphis albella Nevsky, 1951]
Aphis craccivora, fabae, gossypii,
spiraecola
Brachyunguis harmalae
Macrosiphum euphorbiae
Myzus ornatus, persicae
Smynthurodes betae
Toxoptera aurantii

Key to aphids on Hibiscus

1 ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
200 Hippophae

2 Antennae with numerous long hairs. ANT II much longer than ANT I.
ANT PT/BASE less than 0.2. SIPH absent. CAUDA broadly rounded
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smynthurodes betae
- Antennae with sparse, short hairs. ANT II similar in length to ANT I.
ANT PT/BASE more than 0.5. SIPH present. CAUDA tongue-shaped
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyunguis harmalae
3 SIPH long, thin and pale, about 3 x CAUDA and more than 0.3 x BL
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon gossypii
SIPH pale or dark but not more than 2.5 x CAUDA or more than
0.25 x BL . . . . . . . . . . . . . . . . . . see key to polyphagous aphids, p. 532

HIPPOPHAE Sea Buckthorn Elaeagnaceae

All the aphid species found on Hippophae are also recorded from Elaeagnus,
so the key to Elaeagnus aphids can be used to discriminate the species listed
below. It is likely that all heteroecious Capitophorus species can utilize either
Elaeagnus or Hippophae as primary host, depending on availabity.

Host Plant List

Hippophae rhamnoides Capitophorus elaeagni,
hippophaes, meghalayensis,
pakansus, similis, xanthii
Lachnus wichmanni
H. salicifolia Capitophorus similis
Hippophae sp. Capitophorus himalayensis

HOLMSKIOLDIA Verbenaceae

Holmskioldia sp. Aphis gossypii

HOVENIA Rhamnaceae

Hovenia dulcis Myzus persicae

 Ilex 201

HYMENOCARDIA Euphorbiaceae

Hymenocardia acida Aphis gossypii

Toxoptera citricidus
(Use key to polyphagous aphids, p. 532)

HYMENODICTYON Rubiaceae

Hymenodictyon sp. Greenideoidea lambersi

HYMENOSPORUM Pittosporaceae

Hymenosporum flavum Myzus antirrhinii

HYPTIS Labiatae

Hyptis pectinata Sitobion salviae

H. spicata Sitobion salviae
H. suaveolens Sitobion salviae

ILEX Hollies Aquifoliaceae

Host Plant List

Ilex aquifolium Aphis fabae, ilicis
Aulacorthum solani
Illinoia lambersi
Macrosiphum euphorbiae, rosae
202 Ilex

I. cornuta (inch var. burfordii) Macrosiphum euphorbiae

Toxoptera aurantii
I. glabra Macrosiphum rosae
I. integra Macrosiphum rosae
I. macropoda Aulacorthum magnoliae
I. opaca Toxoptera aurantii
I. pedunculosa Toxoptera aurantii, odinae
I. rotunda Myzus persicae
I. serrata Aulacorthum magnoliae
Ryoichitakahashia prunifoltae

Key to aphids on Ilex

1 CAUDA pale, shorter than its basal width, less than 0.3 of length of black
SIPH. Antennal hairs long and fine, mostly 3-4 x basal diameter of ANT
III. ANT III of aptera with secondary rhinaria on distal 0.5-0.7 . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ryoichitakahashia prunifoltae
- CAUDA pale or dark, longer than its basal width, more than 0.3 of length
of SIPH. Antennal hairs less than 3 x basal diameter of ANT III, which
in aptera has rhinaria either absent or only on basal 0.3 . . . . . . . . . . 2
2 Body broadly spindle-shaped. SIPH with an apical zone of polygonal
reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Body oval. SIPH without polygonal reticulation . . . . . . . . . . . . . . . . . . 5
3 SIPH clearly swollen on distal half, proximad to the reticulated part.
Longest hairs on ANT III less than 0.5 x basal diameter of segment
................................................ Illinoia lambersi
- SIPH cylindrical, tapering, or very slightly swollen on distal half.
Longest hairs on ANT III more than 0.5 x basal diameter of segment
............................................................. 4
4 Head black, like SIPH. Dark sclerites present anteriad and posteriad to
SIPH, the antesiphuncular sclerite being crescent-shaped . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum rosae
- Head and SIPH pale or dusky. No dark ante- and postsiphuncular
sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
5 Antennal tubercles well developed, their inner faces parallel or convergent
in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Antennal tubercles poorly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
6 Legs of aptera black except at bases of femora . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
 Inga 203

- Legs of aptera pale except for tarsi and apices of tibiae . . . . . . . . . . . 7

7 Antennal tubercles with inner faces parallel in dorsal view (Fig. 42J). ANT
III of aptera with a small secondary rhinarium near base. SIPH tapering
from base to flange . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- Antennal tubercles with inner faces convergent in dorsal view (e.g.
Fig. 42J). ANT III of aptera never with a secondary rhinarium. SIPH
slightly swollen on distal half . . . . . . . . . . . . . . . . . . . . . Myzus persicae
8 SIPH 0.4-0.6 x CAUDA. ABD TERG 8 with 5-12 hairs . . . . . . . . . . .
............................................... Toxoptera odinae
- SIPH 0.9-1.7 x CAUDA. ABD TERG 8 with 2-5 hairs . . . . . . . . . . . 6
9 Stridulatory ridges present on abdominal sternites 5 and 6, and evenly
spaced Stridulatory peg-like hairs on hind tibiae (Fig. 122). ANT PT/
BASE 3.5-5.0. Dorsal abdominal markings if present confined to ABD
TERG 7 and 8. Alata with media of forewing once-branched . . . . . . .
.............................................. Toxoptera aurantii
- Stridulatory apparatus absent. ANT PT/BASE 2.1-3.2. Dark, broken
sclerites or bands usually present on ABD TERG 1-8. Alata with media
of forewing twice-branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Longest hair on ANT III usually more than 2 x basal diameter of seg-
ment. ABD TERG 2-5 as well as 1 and 7 often with marginal tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis ilicis
- Longest hair on ANT III rarely more than 2 x basal diameter of segment.
ABD TERG 2-5 rarely with small marginal tubercles ... Aphis fabae

INGA Leguminosae

Inga edulis Toxoptera aurantii

I. laurina Sitobion salviae
I. vera Sitobion salviae

SIPH with a subapical zone of polygonal reticulation. No stridulatory

apparatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion salviae
SIPH without polygonal reticulation. Stridulatory apparatus present
(Fig. 122) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
(or try key to polyphagous aphids, p. 532)
204 Iozoste

IOZOSTE = ACTINODAPHNE

JACARANDA Bignoniaceae

Aphis gossypii, spiraecola

Macrosiphum euphorbiae
Myzus ornatus, persicae
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

JUGLANS Walnuts Juglandaceae

According to Bissell (1978), records of Monellia spp. from Juglans may be

considered either as vagrants or as wrongly identified Monelliopsis species.

Host Plant List

Juglans ailanthifolia Dasyaphis rhusae
J. californica Chromaphis californica,
juglandicola
Monelliopsis nigropunctata
J. fallax [Uroleucon fallacis (Nevsky,
1929a)]
J. hindsii Chromaphis juglandicola
J. intermedia = J. regia
J. mandshurica Dasyaphis rhusae
(Garea, Japanese Walnut) Stomaphis asiphon
Toxoptera odinae
J. nigra Chromaphis californica
(American Black Walnut) Monelliopsis caryae,
nigropunctata, tuberculata
J. regia Chromaphis hirsutustibis,
(European Walnut) juglandicola
Panaphis juglandis, nepalensis
Stomaphis mordvilkoi
 Juglans 205

J. ?rupestris Monelliopsis bisetosa, tuberculata

J. sieboldiana Kurisakia ailanthi(?), onigurumi
(Kurumi, Japanese Walnut) [Prociphilus caryae]
Toxoptera odinae
Juglans spp. Aphis spiraecola

Key to aphids on Juglans

1 BL more than 4.5 mm, rostrum longer than body . . . . . . . . . . . . . . . . 2
- BL less than 4.5 mm, rostrum much shorter than body ........... 3
2 SIPH as large pores on broad pigmented hairy cones . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis mordvilkoi
- SIPH absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis asiphon
3 BL more than 3.0mm. Dorsal abdomen with rather broad dark cross
bands on most tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- BL less than 3.0mm. Dorsal abdomen without broad dark cross bands
............................................................. 6
4 Knob of CAUDA globular, about as long as its greatest width (Fig. 52B).
ABD TERG 1 and 2 with broken pigmentation . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panaphis juglandis
- Knob of CAUDA elongate, about 2 x its greatest width (Fig. 52A). ABD
TERG 1 and 2 with solid transverse dark bands . . . . . . . . . . . . . . . . . 5
5 Wing veins often dark but not (or only very weakly) bordered with
fuscous. Pronotum wholly dark. ANT III with 16-26 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panaphis nepalensis nepalensis
- Wing veins bordered with fuscous. Pronotum with a pale central area.
ANT III with 41-50 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panaphis nepalensis yunlongensis
6 Antenna 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Antenna 3- to 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
7 CAUDA dark, tongue-shaped. ANT PT/BASE more than 1.5. Adult
viviparae in colonies mainly apterous . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- CAUDA pale or dusky, knobbed. ANT PT/BASE less than 1.5. Adult
viviparae all alate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 SIPH 0.4-0.6 x CAUDA, which has no constriction (Fig. 121O) . . . . .
............................................... Toxoptera odinae
- SIPH 0.8-1.7 x CAUDA, which has a slight constriction near its mid-
point (Fig. 121T) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
206 Juglans

Fig. 52. CAUDA and anal plate of A, Panaphis nepalensis nepalensis and B, P. juglandis; anal plates of
C, Chromaphis hirsutistibus and D, Ch. californica; E, hairs on middle section of hind tibia of
Ch. hirsutistibus; F, dorsal process of Dasyaphis onigurumi (aptera); G, SIPH of aptera of Kurisakia
onigurumi.

9 ANT PT/BASE less than 0.5. Anal plate only shallowly indented
(Fig. 52C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- ANT PT/BASE more than 0.5. Anal plate deeply bilobed (e.g. Fig. 52D)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Hairs on hind tibia very long, up to about 3.5 x middle diameter of the
tibia (Fig. 52E). Wings with dark spots at ends of veins (use hand lens)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chromaphis hirsutustibis
- Hairs on hind tibia shorter, the longest of them less than 2 x middle
diameter of tibia. Wings without dark spots at ends of veins . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chromaphis juglandicola
11 Forewing veins Cu1a and Cu1b both bordered with fuscous, much darker
than other veins. Each antennal segment darkening over distal half of its
length. Tibiae dark . . . . . . . . . . . . . . . . . . . . . . . . Chromaphis californica
 Juniperus 207

- Forewing veins Cula and Cu1b not darker than other veins, or only
a little darker. Each antennal segment ringed with black apically. Tibiae
pale or dusky . . . . . . . . . . . . . . . . . . Monelliopsis spp. (bisetosa, caryae,
nigropunctata, tuberculata) - use couplets 7-10 of key to aphids on Carya.
12 CAUDA knobbed. Dorsum of aptera bearing numerous long processes,
each tipped with a single bristle (Fig. 52F), these processes as long as or
longer than the reduced, apparently 3-segmented antennae. SIPH as
small, inconspicuous rings . . . . . . . . . . . . . . . . . . . . . . . Dasyaphis rhusae
- CAUDA rounded. Dorsum of aptera without processes, but with numer-
ous long fine hairs. SIPH on broad cones ringed with long hairs (Fig. 52G).
Antennae of both alata and aptera 5-segmented . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kurisakia onigurumi

JULBERNARDIA Leguminosae

Julbernardia globiflora Aphis craccivora

JUNIPERUS Junipers Cupressaceae

The aphid fauna of junipers includes at least 17 species of Cinara, several of

which are known only from their original descriptions. An attempt is made
to discriminate these species in the key below, but for some of the species in
the genus Cinara sensu stricto insufficient samples are available to estimate
the range of seasonal variation, so determinations should be treated warily,
especially in the case of species marked *, specimens of which have not been
seen by the authors.

Host Plant List

Juniperus bermudiana Cinara tujafllina
J. chinensis Cinara fresai, juniperi, tujafilina
Gootiella tremulae
Illinoia morrisoni
J. columnaris Illinoia morrisoni
J. communis (incl. var. hibernica, Cinara cognita, fresai,
nana, sibirica) juniperensis, juniperi,
mordvilkoi, rubicunda,
smolandiae, tujafilina
208 Juniperus

Gootiella tremulae
Illinoia morrisoni
J. conferta Cinara fresai
Illinoia morrisoni
J. horizontalis Cinara cupressi, fresai,
manitobensis, petersoni
Gootiella tremulae
Illinoia morrisoni
J. japonica Cinara fresai
J. macrocarpa Cinara cupressi
J. monosperma Cinara tonaluca
Illinoia morrisoni
J. monticola Illinoia morrisoni
J. occidentalis Cinara fresai, rubicunda
Illinoia morrisoni
J. osteosperma Cinara burrilli, pulverulens,
wahhaca
J. oxycedrus Cinara cupressi, fresai, juniperi,
?petersoni
J. pseudosabina Cinara dahurica, tujafilina
J. sabina [Aploneura juniperina = Gootiella
alba?]
Cinara fresai
Illinoia morrisoni
J. scopulorum Cinara burrilli, cupressi, fresai,
pulverulens, wahluca
Illinoia morrisoni
Sanbornia juniperi
Siphonatrophia cupressi
J. squamata Cinara fresai
Illinoia morrisoni
J. virginiana Cinara cupressi, fresai,
juniperivora
Illinoia morrisoni
Sanbornia juniperi
Siphonatrophia cupressi

Key to juniper aphids (apterae viviparae only)

1 SIPH long and tubular, swollen on distal half, with subapical reticulation
(Fig. 53A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni
 Juniperus 209

- SIPH (if visible) not long and tubular . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 SIPH absent, or inconspicuous pores, or small pale cones . . . . . . . . . 3
- SIPH in form of large pores on broad, often pigmented, hairy cones
............................................................. 5
3 CAUDA longer than its basal width. ANT PT/BASE more than 0.5. Wax
pore plates not developed. Legs normal . . . . . . . . . . . . . . . . . . . . . . . . . 4
- CAUDA broadly rounded, much shorter than its basal width. ANT PT/
BASE less than 0.25. Wax pore plates on posterior dorsal abdomen. Legs
very thick, and tibiae armed with spines distally (Fig. 53B) . . . . . . . . . .
.............................................. Gootiella tremulae
4 Antenna usually 4-segmented. Frons with a median quadrate projection
(Fig. 53C). SIPH reduced to minute pores hardly larger than spiracles.
CAUDA long and tapering, with more than 20 hairs . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sanbornia juniperi
- Antenna 5- or 6-segmented. Frons without a median projection. SIPH
small, pale cones (Fig. 44A). CAUDA tongue-shaped with about 10 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphonatrophia cupressi
5 Hairs on ANT III sparse and very short, less than 0.5 x diameter of
segment at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara wahhaca
- Hairs on ANT III mostly longer than diameter of segment at midpoint
............................................................. 6
6 Antennae and legs densely clothed in fine, rather short, rather recumbent,
hairs; those on ANT III mostly less than 1.5 x diameter of segment at
midpoint, and those on hind tibia mostly less than width of tibia at mid-
point (Fig. 53D). SIPH base with more than 70 very fine hairs, shorter
than diameter of siphuncular pore . . . . . . . . . . . . . . Cinara juniperivora
- Hairs less fine and numerous, more erect and usually longer. SIPH base
if with more than 70 hairs then many of these are longer than diameter
of siphuncular pore . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Hairs on ANT III mostly less than 2 x diameter of segment at midpoint.
Hairs on dorsal side of hind tibia usually less than 1.5 x width of tibia
at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Hairs on ANT III mostly more than 2 x diameter of segment at midpoint.
Hairs on dorsal side of hind tibia usually more than 1.5 x width of tibia
at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
8 SIPH cones large, of maximum basal diameter more than 0.3mm
.............................................. Cinara rubicunda*
- SIPH cones of maximum basal diameter less than 0.27 mm . . . . . . . . 9
9 ANT PT/BASE more than 0.5. Diameter of base of SIPH cone less than
0.12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara tonaluca*
210 Juniperus

Fig. 53. A, SIPH of Illinoia morrisoni (aptera); B, hind leg of aptera of Gootiella tremulae; C, ventral
aspect of head of Sanbornia juniperi aptera; D, middle section of hind tibia of Cinara juniperivora
(aptera); E, hind tarsus of Cinara wahluca showing measurement of dorsal length and basal width;
F, same for C. juniperi; G, ANT VI of C. juniperi; H, ANT VI of C. petersoni.
 Juniperus 211

- ANT PT/BASE less than 0.4. Diameter of base of SIPH cone more than
0.14 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 ANT BASE VI less than 0.5 x HT II ... Cinara burrilli*/pulverulens
- ANT BASE VI more than 0.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Dorsum with numerous hairs, not confined to transverse rows. ANT III
with a rhinarium near apex . . . . . . . . . . . . . . . . . . . Cinara manitobensis
- Dorsum with fewer hairs arranged in transverse rows. ANT III usually
without a primary rhinarium . . . . . . . . . . . . . . . . . . . . . . Cinara cognita*
12 HT I with dorsal length a little more than, or at least equal to (1.0-1.2 x)
basal width (e.g. Fig. 53E). PT with 4-5 subapical hairs . . . . . . . . . . 13
- HT I with dorsal length distinctly shorter than (0.5-0.8 x) basal width
(e.g. Fig. 53F). PT usually with 3 subapical hairs (subgenus Cupressobium)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13 Greatest diameter of SIPH cone 0.25 mm, less than R IV+V. HT II less
than 0.25 mm. Base VI less than 0.14mm . . . . . . . . . . Cinara wahluca
- Greatest diameter of SIPH cone at least 0.28mm, more than R IV+V.
HT II more than 0.25mm long. Base VI usually more than 0.14mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara juniperensis
14 Femora usually wholly pale except at point of articulation with tibia,
and tibiae only dark at apices. ANT PT/BASE usually about 0.2. R IV
0.14-0.18 mm long, HT II 0.20-0.28 mm long . . . . . . Cinara tujafilina
- Femora distally dark and tibiae either wholly dark or dark at base as well
as apex. If ANT PT/BASE is less than 0.25 then R I V is more than 0.2 mm
long and HT II is more than 0.3 mm long . . . . . . . . . . : . . . . . . . . . . . 15
15 ANT VI BASE with 4-7 hairs restricted to basal half (Fig. 44B). Tibiae
only dark at base and apex . . . . . . . . . . . . . . . . . . . . . . . Cinara cupressi
- ANT VI BASE with 7-15 hairs not restricted to basal half. Tibiae wholly
dark or paler in middle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 BL more than 3.3 mm. Length of sclerotized part of stylet groove (Fig. 74A,
p. 292) more than 1.5 mm, and hind tibia more than 2.2 mm long. Hairs on
hind tibia about as long as its diameter at midpoint . . . . . . . . . . . . . . . 17
- BL usually less than 3.3 mm, rarely up to 3.5 mm. Length of sclerotized
part of stylet groove less than 1.2mm, length of hind tibia less than
2.15mm, with dorsal tibial hairs at least 1.5 x width of hind tibia at
midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 ANT PT less than 0.05mm, ANT PT/BASE 0.17-0.22. R IV+V
0.35-0.4mm, 4.8-5.0 x its basal width . . . . . . . . . . . . Cinara dahurica*
- ANT PT more than 0.05mm, ANT PT/BASE 0.22-0.33. R IV+V
0.29-0.34 mm, 3.1-4.1 x its basal width . . . . . . . . . Cinara smolandiae
212 Kalopanax

18 Hind tibiae with pale section on basal half. ANT III usually longer than
maximum diameter of base of SIPH cone . . . . . . . . . . . . Cinara fresai
- Hind tibiae wholly dark. ANT III usually shorter than maximum diameter
of base of SIPH cone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 ANT BASE VI narrow and often slightly constricted at about its midpoint
(Fig. 53G); width at midpoint usually less than 0.2 of length (look at
several specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara juniperi
- ANT BASE VI broader, width at midpoint more than 0.2 of length (e.g.
Fig.53H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 ANT BASE VI longer than both ANT IV and R IV . . . . . . . . . . . . . . .
................................................ Cinara petersoni
- ANT BASE VI shorter than both ANT IV and R IV . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara mordvilkoi

KALOPANAX Araliaceae

Kalopanax ricinifolius Aphis kalopanacis

KETELEERIA Pinaceae

Keteleeria evelyniniana Mindarus keteleerifoliae

K. fortunei Cinara keteleeriae

Densely hairy aphid, BL more than 3 mm. Aptera with multifaceted eyes.
Secondary rhinaria rounded. SIPH as large pores on shallow pigmented
hairy cones. CAUDA rounded with numerous hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara keteleeriae
Sparsely haired aphid, BL less than 2.5 mm. Aptera with 3-faceted eyes.
Secondary rhinaria of alata transversely elongate. SIPH as small pores
without surrounding hairs or pigmentation. CAUDA triangular with 2
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mindarus keteleerifoliae
 Laburnum 213

KHAYA African Mahoganies Meliaceae

Khaya senegalensis Aphis gossypii

KIELMEYERA Guttiferae

Kielmeyera coriacea Aphis gossypii

KIGELIA Bignoniaceae

Kigelia africana Aphis gossypii

Toxoptera aurantii
K. pinnata Aphis gossypii
(See key to polyphagous aphids, p. 532)

KOELREUTERIA Sapindaceae

Koelreuteria paniculata Periphyllus koelreuteriae

KRAUSSIA Rubiaceae

Kraussia floribunda Aphis spiraecola

LABURNUM Leguminosae

Laburnum anagyroides Aphis craccivora, cytisorum

Myzus persicae
214 Laccosperma

Key to aphids on Laburnum

1 Antennal tubercles well developed, convergent in dorsal view (Fig. 42I).
SIPH slightly swollen on distal half. SIPH and CAUDA pale . . . . . . .
................................................. Myzus persicae
Antennal tubercles poorly developed. SIPH tapering. SIPH and CAUDA
dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 BL divided by R IV+V less than 3.2+ (6 x BL) . . . . . Aphis cytisorum
- BL divided by R IV+V more than 3.2 +(6 x BL) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora

LACCOSPERMA Palmae

Laccosperma sp. Cerataphis variabilis

(For aphids on palms see B & E, 1984.)

LAGERSTROEMIA Crape-Myrtles Lythraceae

Lagerstroemia flos-reginae Aphis gossypii

Tinocallis (Sarucallis) himalayensis
L. indica Aphis gossypii, spiraecola
Aulacorthum magnoliae
Myzus persicae
Tinocallis (Sarucallis)
kahawaluokalani
Toxoptera aurantii
L. macrocarpa Tinocallis (Sarucallis) khonkaensis
Lagerstroemia sp. Tinocallis (Sarucallis) khonkaensis

Key to Lagerstroemia aphids

1 All adult viviparae winged. CAUDA knobbed, anal plate bilobed.
ANT PT/BASE less than 1.5. SIPH as short, pale, truncated cones
............................................................. 2
Adult viviparae apterous or alate. CAUDA tongue-shaped, anal plate
entire. ANT PT/BASE more than 1.5. SIPH tubular, tapering or swollen
on distal half . . . . . . . . . . . . . . . use key to polyphagous aphids, p. 532
 Lansium 215

Fig. 54. Spinal tubercles on ABD TERG 2 of A, Tinocallis kahawaluokalani and B, T. himalayensis.

2 Forewings marked with broad bands of fuscous, especially along and

behind the main vein and along the branches of the media. Head and
thorax without spinal tubercles. Spinal tubercles on ABD TERG 2 large,
dark, united over more than half their length (Fig. 54A) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis (Sarucallis) kahawaluokalani
Forewings with veins often lightly bordered but without broad bands of
fuscous. Head and thorax with spinal tubercles. Spinal tubercles on ABD
TERG 2 pale or dusky, separate or fused only at bases (Fig. 54B) . . .
. . . . . . . . . Tinocallis (Sarucallis) himalayensis/khonkaensis (see p. 904)

LANNEA Anacardiaceae

Lannea acidissima Toxoptera aurantii

L. edulis Aphis gossypii
L. ?stuhlmanni Aphis gossypii
L. wodier Toxoptera odinae
(Use key to polyphagous aphids, p. 532)

LANSIUM Meliaceae

Lansium ?domesticum Schizoneuraphis gallarum

216 Laportea

LAPORTEA Urticaceae

Laportea aestuans Sitobion autriquei

L. bulbifera Hydronaphis laporteae

Key to aphids on Laportea

SIPH black, slightly tapering and less than 2 x CAUDA. Body and
antennae bearing hairs shorter than diameter of ANT III . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion autriquei
SIPH dusky, weakly clavate with a well-developed flange, more than
3 x CAUDA. Hairs on body and antennae much longer than diameter of
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydronaphis laporteae

LARIX Larches Pinaceae

The aphid fauna of Larix seems to reflect the apparently close relation-
ship between the tree species in this genus. Host records of the best-known
larch-feeding aphids (Cinara cuneomaculata, C. laricis, Adelges laricis)
show no sign of discrimination between species of Larix, and it seems possible
that the less well-known and less widely-distributed aphids would be equally
non-specific in their choice of hosts if given the opportunity. For Adelges
species, it is not possible to provide a complete key in the present state of
knowledge.

Host Plant List

Larix cajanderi Cinara laricis
L. decidua Adelges diversis, lands, segregis,
(European Larch) viridana, viridis
[Aphis fabae group]
Cinara cuneomaculata, kochiana,
laridcola, larids
L. x eurolepis Adelges viridana, viridis
Cinara larids
L. gmelinii Adelges karamatsui, larids,
(Dahurian Larch) viridana
Cinara cuneomaculata, kochiana,
laridcola, larids
 Larix 217

L. kaempferi = L. leptolepis
L. kamtschatica Cinara cuneomaculata, kochiana,
laricicola, laricis
L. koraiensis (koreana?) Adeiges karamatsui
Cinara kochiana ssp. kochi
L. laricina Adelges aenigmaticus, lariciatus,
(Alaskan Larch, Tamarack) laricis
Cinara cuneomaculata, laricifex,
spiculosa, subterranea
L. leptolepis Adelges isedakii, karamatsui,
(Japanese Larch, Karamatsu) kitamiensis, laricis, torii,
viridana
Cinara chibi, cuneomaculata,
kochiana, kochiana ssp. kochi,
laricicola, laricis
Prociphilus [?kuwanai], laricis,
[ushikoroshi]
L. lyalli Adelges lariciatus
Cinara lyalli
L. occidentalis Adelges oregonensis
(Western Larch) Cinara laricifoliae
L. pontaninii Adelges laricis ssp. pontaninilaricis
L. sibirica Adelges tardoides, viridana,
(Siberian Larch) viridis, viridula
Cinara cuneomaculata
Elatobium laricis

Key to larch aphids

1 Antennae of aptera with at most 3 segments. Antenna of alata 4- or
5-segmented with 3 large primary, but no secondary, rhinaria. Forewing
with Rs absent. Always oviparous, with a distinct, chitinous ovipositor
............................................................. 2
- Antenna of aptera and alata 6-segmented, that of alata with secondary
rhinaria. Forewing with Rs present. Parthenogenetic morphs viviparous
............................................................. 4
2 Wax glands of overwintering first instar larva (neosistens) with double-
walled facets secreting hollow tubes of wax (Fig. 55A). Head and pro-
thorax of alata bearing wax gland areas of granular appearance, with
facets not clearer than the surrounding cuticle, or at most with small
groups of clear facets on marginal areas of prothorax (Fig. 55D) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges (Sacciphantes) viridis group
218 Larix
 Larix 219

- Wax glands of overwintering neosistens either absent or if present then

with simple, thin-walled facets secreting solid wax filaments (Fig. 55B).
Head and prothorax of alata with or without wax gland areas bearing
numerous facets that are clearer than surrounding cuticle (Figs 55E,F)
............................................................. 3
3 Overwintering first instar larva (neosistens) with dorsal sclerotic plates of
head and thorax separate (Fig. 55B). Adult aptera (sistens) pyriform, BL
1.8-2.5 mm. Dorsum membranous, with small groups of wax gland facets
on all segments (the marginal groups largest) (Fig. 55G). Alata of BL
1.8-2.6 mm, with anterior and posterior cephalic wax gland areas united
to form a continuous longitudinal swathe of facets on each side of the
epicranial suture (Fig. 55E). ANT IV and V of alata with minimal width
of basal articulation of segment usually less than 0.5 of width of rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . Adelges (Cholodkovskyana) viridana group
- Overwintering neosistens heavily sclerotized, with dorsal sclerotic plates
of head and thorax coalesced (Fig. 55C). Adult aptera (sistens or pro-
grediens) oval or subcircular, with paired spinal, pleural and marginal
sclerotic plates on thorax and ABD TERG 1-5; these are rounded, convex
and wrinkled without wax glands in sistens (spring generation; Fig. 55H),
but more extensive and with numerous wax gland facets in aestivating
(progrediens) generation (Fig. 55I). Alata of BL 1.0-1.7 mm, with anterior
and posterior cephalic wax gland areas if present usually separate
(Fig. 55F). ANT IV and V of alata with minimum width of basal articula-
tion of segment usually more than 0.5 of maximum width of rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges laricis group
4 SIPH tubular, longer than their basal width. ANT PT/BASE more than
1 ............................................................ 5
- SIPH either absent or on broad, hairy, pigmented cones. ANT PT/BASE
less than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
5 Dorsum pale. CAUDA pale. SIPH mainly pale, and swollen on distal
half. Antennal hairs very short and inconspicuous, of maximal length less
than 0.5 x basal diameter of ANT III . . . . . . . . . . . . Elatobium laricis
- Dorsum with dark markings, at least marginally and posterior to SIPH.
CAUDA dark. SIPH dark, tapering from base to flange. Antennal hairs
more than 0.5 x basal diameter of ANT III . . . . . . . . . . . Aphis fabae
6 SIPH absent or present only as small pores . . . . . . . . . . . . . . . . . . . . . 7

Fig. 55. A, head of neosistens of Adelges viridis showing wax glands with double-walled facets; head
and prothorax of B, A. viridana neosistens and C, A. laricis (partly after Schneider-Orelli and Schneider,
1954); head and prothorax of alata (sexupara) showing arrangement and details of wax glands of
D, A. viridis, E, A. viridana and F, A. laricis; G, adult sistens of A. viridana (in May); H, adult sistens
of A. laricis (in April); I, adult progrediens of A. laricis (July).
220 Larix

Fig. 56. Sclerites on ABD TERG IV of apterae of A, Cinara lyalli, B, C. laricifoliae, C, C. laricifex,
D, C. spiculosa, E, C. laricis.

- SIPH present as large pores on broad hairy cones . . . . . . . . . . . . . . . . 9

7 Alata developing on Larix (sexupara) with only about 6 secondary rhinaria
on ANT III, 2-4 on ANT IV, about 3 on V, and 2 on VI . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus laricis
- Alate sexupara with many more secondary rhinaria . . . . . . . . . . . . . . . 8
8 Head of alata without distinct wax pore plates (based on spring migrant,
but probably also applying to sexupara) . . . . . . . . Prociphilus kuwanai
- Head of alata with a pair of wax pore plates (based on spring migrant,
but probably applying also to sexupara) . . . . . Prociphilus ushikoroshi
9 (This and subsequent couplets apply only to APTEROUS morphs.) ABD
TERG 2-6 with at least some hairs arising from dark sclerites of minimum
dimension at least 5 x the diameter of the hair base . . . . . . . . . . . . . 10
- ABD TERG 2-6 either without hair-bearing sclerites or with hairs on
very small, round sclerites of diameter less than 4 x that of hair base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
10 ANT BASE VI with 9-12 hairs. Hair-bearing sclerites on ABD TERG
2-6 numerous rounded, evenly-spaced, and mostly rather uniform-sized
(Fig. 56A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara lyalli
- ANT BASE VI with 6-8 hairs. Hair-bearing sclerites on ABD TERG 2-6
of irregular shape and varying size and distribution . . . . . . . . . . . . . 11
11 Hind tibia mainly dark with a short pale section on basal third, of length
less than 0.2 of total length of tibia. Hairs on hind tibia maximally 65 µm
 Larix 221

and hair-bearing sclerites on ABD TERG 2-6 numerous, mostly separate

(Fig. 56B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara laricifoliae
- Hind tibia with most of basal half pale, the pale section occupying at least
0.25 of total length of tibia, if the longest hind tibial hairs are less than
65 µm then there are only a few, small sclerites on each of ABD TERG
2-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 R IV+V usually less than 0.7 times longer than HT II. ABD TERG 2-6
each with only a few small flat sclerites, bearing variable but often rather
finely pointed hairs which are maximally 40 µm long (Fig. 56C) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara laricifex
- R IV+V usually more than 0.7 times longer than HT II. ABD TERG 2-6
usually with numerous large sclerites, often confluent between hair bases;
the hairs arising from them are often thick and spine-like with raised,
conical bases (Figs 56D, E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Dorsal abdominal hairs often with enlarged or blunt, bifurcate or multi-
furcate apices (Fig. 56D) . . . . . . . . . . . . . . . . . . . . . . . . . Cinara spiculosa
- All dorsal abdominal hairs with acute apices (Fig. 56E) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara laricis
14 BL more than 4.8mm. R IV 0.29-0.42 mm long, bearing 21-34 hairs
arranged in 4 longitudinal rows . . . . . . . . . . . . . . . . . . . Cinara kochiana
- BL less than 4.8mm. R IV 0.15-0.25 mm long, bearing 5-11 hairs in 2
longitudinal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Longest hairs on ABD TERG 3 at least 2 x diameter of ANT III at
midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- Longest hairs on ABD TERG 3 no more than 1.5 x diameter of ANT III
at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 ANT VI (BASE + PT) less than 0.16mm long, distinctly shorter than R
IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara laricina
- ANT VI (BASE + PT) more than 0.2 mm long, similar in length to or
slightly longer than R IV . . . . . . . . . . . . . . . . . . . . . Cinara subterranea*
17 ANT IV shorter than or as short as ANT VI (incl. PT). Length of
sclerotized part of stylet groove (see Fig. 74A) more than total length of
antenna. BL usually less than 3 mm . . . . . . . . . . . . . . . . . . . Cinara chibi
- ANT IV clearly longer than ANT VI (incl. PT). Length of sclerotized part
of stylet groove less than total length of antenna. BL often more than
3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara cuneomaculata/laricicola

*0nly oviparae of C. subterranea have been seen.

222 Laurus

LAURUS Lauraceae

Laurus rotundifolia Aphis spiraecola

LEPTOSPERMUM Myrtaceae

Leptospermum laevigatae Myzus ornatus

L. scoparium Myzus persicae
Leptospermum sp. Anomalaphis casimiri

ABD TERG 7 and 8 each bearing a pair of long, backwardly directed

processes, those on ABD TERG 7 about as long as SIPH, which have a
subapical zone of reticulation and a ring of 4-5 hairs with furcate apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anomalaphis casimiri
ABD TERG 7 and 8 without long processes. SIPH without reticulation
and hairs . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

LEUCADENDRON Proteaceae

Leucadendron argenteum Aphis gossypii

LEUCAENA Leguminosae

Leucaena glauca Aphis fabae group

LEUCOSIDEA Rosaceae

Leucosidea sericea Aphis gossypii

 Ligustrum 223

LIGUSTRUM Privets Oleaceae

Host Plant List

Ligustrum folicosum Aphis crinosa
L. japonicum Aphis crinosa
Prociphilus ligustrifoliae
L. obtusifolium ( = ibota) Aphis crinosa
Aphis spiraecola
Aulacorthum ibotum
Prociphilus oriens
L. ovalifolium Myzus ligustri
Prociphilus oriens
L. sinense [Chaitoregma sp.? (galls)]
L. tschonoskii var. glabrescens Prociphilus oriens
L. vulgare Aphis gossypii
Macrosiphum euphorbiae
Myzus ligustri, persicae
Prociphilus bumeliae
Ligustrum sp. [Prociphilus cheni]
(Note: some records from the common European privet L. vulgare may be
erroneous, due to misidentification of the planted hedge privet L. ovalifolium.)

Key to aphids on Ligustrum

1 SIPH absent, ANT PT/BASE less than 0.25 . . . . . . . . . . . . . . . . . . . . . 2
- SIPH present, ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . 4
2 ANT VI BASE of alata (from leaf-nest gall) with 0-7 secondary rhinaria.
Forewing veins bordered with fuscous . . . . . . Prociphilus ligustrifoliae
ANT VI BASE of alata (from leaf-nest gall) always without any secondary
rhinaria. Forewing veins not fuscous-bordered . . . . . . . . . . . . . . . . . . . 3
3 ANT V of alata (from leaf-nest gall) with 7-10 secondary rhinaria . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
ANT V of alata (from leaf-nest gall) with 0-2 secondary rhinaria . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus bumeliae
4 Longest hairs on ANT III more than 3 x basal diameter of segment. SIPH
short, truncate, similar in length to the short CAUDA, which bears more
than 14 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis crinosa
224 Lindera

Longest hairs on ANT III less than 2 x (and usually shorter than) basal
diameter of segment. SIPH usually clearly longer than CAUDA, which
bears less than 14 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 SIPH conspicuously swollen over about 0.7 of length, and in aptera
having basal half pale and distal half dark. Alata with secondary rhinaria
on ANT IV and a black dorsal abdominal patch . . . . . Myzus ligustri
SIPH tapering or slightly swollen on distal half, but if swollen then in
aptera only dark at extreme apices. Alata without secondary rhinaria on
ANT IV, with or without a black dorsal patch . . . . . . . . . . . . . . . . . . . 6
6 SIPH long, more than 0.2 x BL, and uniformly dark in aptera, much
darker than CAUDA; without any polygonal reticulation . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum ibotum
SIPH either much less than 0.2 x BL or if longer then not much darker
than CAUDA in aptera, and with a subapical zone of polygonal reticula-
tion . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

LINDERA Lauraceae

Host Plant List

Lindera erythrocarpa Aulacorthum muradachi
L. glauca Aiceona actinodaphinis
L. praecox Aulacorthum muradachi
Thoracaphis linderae
L. pulcherrima Aiceona parvicornis
L. sericea Aulacorthum linderae
L. triloba Aulacorthum muradachi
Thoracaphis linderae
L. umbellata Aulacorthum muradachi
Thoracaphis linderae
Lindera sp. ?Metanipponaphis sylvestrii
[Paratrichosiphum sp. - David
et al., 1969]

Key to aphids on Lindera

1 Aptera aleyrodiform; body flattened, almost circular, with dark sclerotic
dorsum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Aptera of normal aphid form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
 Liquidambar 225

2 Marginal hairs long and stout . . . . . . . . . . . . . . . . Thoracaphis linderae

Marginal hairs fine . . . . . . . . . . . . . . . . . . . . ?Metanipponaphis sylvestrii
3 SIPH pores placed on hairy cones. ANT PT/BASE less than 1 . . . . 4
- SIPH tubular. ANT PT/BASE more than 2 . . . . . . . . . . . . . . . . . . . . . 5
4 R IV+V 0.76-0.80 x HT II and bearing 2-3 accessory hairs. SIPH cones
small, basal diameter maximally 0.12mm . . . . . . . Aiceona parvicornis
R IV+V 1.2-1.4 x HT II and bearing 6 or more accessory hairs. SIPH
cones large, of basal diameter more than 0.2 mm . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona actinodaphnis
5 SIPH with numerous long hairs . . . . . . . . . . . . . . Paratrichosiphum sp.
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
6 SIPH of aptera much darker than CAUDA. R IV+V 0.8-1.0 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum muradachi
SIPH of aptera pale, or dark only at apices. R IV+V 1.2-1.4 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum linderae*

LIQUIDAMBAR Sweet Gums Hamamelidaceae

Liquidambar formosana Aphis gossypii

(Thau) Longistigma liquidambaris
Myzus persicae
Stomaphis liquidambarus
L. orientalis Myzus persicae
L. styraciflua Aphis gossypii
(American Red Gum) Aulacorthum solani
Longistigma liquidambaris
Myzus ornatus

Key to aphids on Liquidambar

1 BL less than 3 mm. CAUDA tongue-shaped . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
BL more than 4 mm. CAUDA broadly rounded . . . . . . . . . . . . . . . . . . 2
2 Rostrum longer than body. Forewing of alata with veins bordered with
fuscous, and with pterostigma not extending to wing-tip . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis liquidambarus

*Not seen by authors.

226 Liriodendron

Rostrum much shorter than body. Forewing of alata with wing veins not
or only very lightly bordered, and with pterostigma extended to wing-tip
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma liquidambarus

LIRIODENDRON Magnoliaceae

Liriodendron tulipifera Aphis fabae

(Tulip Tree) Illinoia liliodendri
Macrosiphum euphorbiae
Myzus persicae

Key to aphids on Liriodendron

1 SIPH without polygonal reticulation. ANT III of aptera without any
secondary rhinaria . . . . . . . . . go to key to polyphagous aphids, p. 532
SIPH with a subapical zone of polygonal reticulation. ANT I I I of aptera
with 2 or more secondary rhinaria on basal half . . . . . . . . . . . . . . . . . 2
2 SIPH 2.4-2.8 x CAUDA. SIPH of aptera clearly darker than CAUDA
and usually somewhat swollen on distal half . . . . . . Illinoia liriodendri
SIPH 1.7-2.3 x CAUDA; in aptera not darker than CAUDA except at
apices, and cylindrical on distal half . . . . . . . Macrosiphum euphorbiae

LITHOCARPUS (= PASANIA) Fagaceae

Host Plant List

Lithocarpus bennettii [Eutrichosiphum sp. (Noordam, in
prep.)]
[Greenidea sp. (Noordam, in
prep.)]
[Mesotrichosiphum sp. (Noordam,
in prep.)]
Nipponaphis semiglabra
L. cuspidata Eutrichosiphum pasaniae
Mollitrichosiphum tenuicorpus
[Myzocallis punctata]
L. densiflora Tuberculatus (Pacificallis) pasaniae
L. edulis Greenidea (Trichosiphum) kuwanai
 Lithocarpus 227

L. glabra Eutrichosiphum dubium,

heterotrichum
Greenidea (Trichosiphum) kuwanai
Mollitrichosiphum lithocarpi
L. indutus Sinonipponaphis hispida
L. konishii Eutrichosiphum dubium
L. rhombocarpa Greenidea (Trichosiphum) nigra
L. sundaicus Metanipponaphis vandergooti
L. uraiana Mollitrichosiphum lithocarpi
Lithocarpus spp. Dermaphis japonensis, takahashii
[Eutrichosiphum flavum
Takahashi]
Lithoaphis lithocarpi
Metanipponaphis lithocarpicola
[Mollitrichosiphum nigrofasciatum]
Parathoracaphis setigera
Uichancoella gabrieli

Key to aphids on Lithocarpus

1 Body of aptera aleyrodiform; oval, strongly sclerotized dorsally with
head, thorax and ABD TERG 1 more-or-less fused (as prosoma). Legs and
antennae much reduced and SIPH pore-like or absent. Antennae of alata
with annular rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Aptera of normal aphid form with well-developed legs and antennae.
SIPH either truncated cones or tubes clothed with long hairs. Antennae
of alata with oval or transversely elongate rhinaria . . . . . . . . . . . . . . 10
2 Aptera without SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Aptera with SIPH as a pair of pores (sometimes very small) on abdominal
plate (fused ABD TERG 2-7; see Fig. 36B) . . . . . . . . . . . . . . . . . . . . . . 5
3 Body of aptera rather narrow, flattened dorsally, with stout, spine-like
marginal hairs (Fig. 91F) . . . . . . . . . . . . . . . . . . Parathoracaphis setigera
- Body of aptera broadly oval, convex dorsally, with thin, pointed marginal
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Marginal hairs of aptera 120-160 µm long (Fig. 91L) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermaphis japonensis
- Marginal hairs of aptera maximally about 100µm long . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermaphis takahashii
5 Prosoma of aptera densely covered with non-articulated hair-like pro-
cesses, 15-40 µm long, lacking only on muscle plates (Fig. 57A) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinonipponaphis hispida
228 Lithocarpus

Fig. 57. A, margin of prosoma of Sinonipponaphis hispida, with hair-like processes; B, prosomal
marginal hairs of Uichancoella gabrielli; C, ventrolateral part of abdomen of Mollitrichosiphum tenuicorpus
and D, ridges on basal part of hind tibia (also present in M. lithocarpi).

- Prosoma of aptera without hair-like processes . . . . . . . . . . . . . . . . . . . 6

6 Prosoma of aptera densely ornamented with numerous separate oval or
rounded papillae or pustules (e.g. Fig. 36C). Marginal hairs numerous,
minute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Prosoma of aptera variably sculptured, but without a dense ornamenta-
tion of separate rounded papillae or pustules. Marginal hairs more than
20 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 Prosoma of aptera with 3-13 hairs 30-50 µm long on medial area of each
segment. Total antennal length 0.13-0.14 x BL . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metanipponaphis vandergooti
- Prosoma of aptera without any hairs on medial area (?). Total antennal
length only about 0.06-0.07 x BL ... Metanipponaphis lithocarpicola
8 Prosoma of aptera fused with abdominal plate (e.g. Fig. 36C). Marginal
hairs short and spine-like, about 28 µm long, and with 5 pairs of very
short spinal hairs . . . . . . . . . . . . . . . . . . . . . . . . . . Lithoaphis lithocarpi*
- Prosoma and abdominal plate at least partially separate. Marginal hairs
not spine-like, 50-180µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Prosoma of aptera with marginal hairs very long, stout basally but with
finely-pointed apices, about 160 µm long (Fig. 57B); with 5 pairs of similar
large spinopleural hairs, the 3 thoracic tergites also bearing several some-
what shorter, thinner hairs . . . . . . . . . . . . . . . . . . . Uichancoella gabrieli
- Prosomal hairs of aptera all 50-80 µm long . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis semiglabra*
 Lithocarpus 229

10 SIPH as truncated cones. CAUDA knobbed, anal plate bilobed. ABD

TERG 1 and 2 each with a pair of large spinal processes. (All viviparae
alate) . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus (Pacificallis) pasaniae
- SIPH as hairy tubes. CAUDA much broader than long, anal plate entire.
ABD TERG 1 and 2 without processes. (Viviparae apterous or alate)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Antennae of both aptera and alata 5-segmented . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pasaniae
- Antennae usually 6-segmented, occasionally 5-segmented in aptera . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Hind tibia with a series of about 15 tranverse (stridulatory) ridges spaced
out over proximal 0.5-0.6 of length (Fig. 57D). Body of aptera about
2.5-3.0 x longer than its maximum width . . . . . . . . . . . . . . . . . . . . . . 13
- Hind tibia without stridulatory ridges. Body of aptera less than 2 x its
maximum width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13 Ventrolateral areas of abdomen densely spinulose (Fig. 57C). ANT III of
aptera at least 1.8 x ANT IV+V together . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum tenuicorpus
- Ventrolateral areas of abdomen not spinulose. ANT III of aptera about
1.5 x ANT IV+V together . . . . . . . . . . . . Mollitrichosiphum lithocarpi*
14 R IV+V very long, 2.5-3.1 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- R IV+V 1.3-1.8 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
15 SIPH of aptera pale, shorter than head width across eyes, with long hairs
all of similar length (except near base) . . . . . . Eutrichosiphum flavum
- SIPH of aptera dark, longer than head width across eyes, with hairs of
very different lengths interspersed, the shortest hairs being about half as
long as the longest (Fig. 92G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 ANT III imbricated (Fig. 92J) . . . . . . . . . . . . . . Eutrichosiphum dubium
- ANT III smooth . . . . . . . . . . . . . . . . . . . . Eutrichosiphum heterotrichum
17 SIPH of alata similar in pigmentation to hind tibiae, 0.40-0.65 x BL and
8.0-12.5 x longer than their width at midpoint . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) kuwanai
- SIPH of alata much darker than hind tibiae, 0.65-0.75 x BL and
16.5-17.5 x their width at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) nigra
230 Litsea

LITSEA Lauraceae

Host Plant List

Litsea amara Pentatrichosiphum luteum
Schizoneuraphis litseicola
L. chinensis Schizoneuraphis gallarum
L. corymbosa Nipponaphis manoji
L. cubeba Eutrichosiphum sankari
L. glauca Nipponaphis litseae
L. monopetala Aiceona titabarensis
L. polyantha Aiceona robustiseta, titabarensis
Greenideoida (Pentatrichosiphum)
lutea
Nipponaphis manoji, Nipponaphis
sp. (Raha and Raychaudhuri,
1981)
Schizoneuraphis himalayensis
Sinomegoura citricola
L, populifolia Aiceona actinodaphnis
L. pungens Aiceona actinodaphnis
L. sebifera Eutrichosiphum dubium, litseae
Hoplothoracaphis manipurense
L. veitchiana Aiceona actinodaphnis
L. wilsonii Aiceona actinodaphnis
Litsea sp. Eutrichosiphum makii

Key to aphids on Litsea

1 Aptera aleyrodiform; dorsum with head, thorax and ABD TERG 1 fused
as prosoma, ABD TERG 2-7 reduced and fused together (as abdominal
plate); and much reduced, 3- or 4-segmented antennae . . . . . . . . . . . . 2
- Aptera of normal aphid form, with 4- to 6-segmented antennae . . . . 7
2 Aptera without SIPH; marginal hairs stout, dagger-like, about 40 µm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplothoracaphis manipurense
- Apterae with SIPH as small pores; marginal hairs variable, not dagger-like
............................................................. 3
 Litsea 231

Fig. 58. Marginal prosomal hairs of apterae of A, Schizoneuraphis gallarum and B, S. litseicola] C, part
of surface of prosoma of Nipponaphis litseae showing pustulate ornamentation and dorsal hairs;
abdominal plate of D, S. himalayensis, E, Schizoneuraphis gallarum and F, S. litseicola.

3 Aptera with large areas of dorsal cuticle of prosoma granular or almost

smooth, regions of pustulate ornamentation being localized. Longest pro-
somal hairs very thick (6-10 µm basal diameter) and more t h a n 100 µm
long, with raised bases; ABD TERG 8 with a crenulate margin . . . . 4
- Aptera with dense pustulate ornamentation extending over most of dorsal
area of prosoma (Fig. 58C). Longest prosomal hairs are fine and much
less than 100µm long, without raised bases. ABD TERG 8 without a
crenulate margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4 Aptera with prosomal hairs maximally 140 µm long. Abdominal plate with
the first 3 pairs of submarginal hairs long, stout and dark, much larger
and thicker (e.g. basal diameter 3-4 x more) than the posterior 3 pairs
(Fig. 58D) . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneuraphis himalayensis
- Aptera with prosomal hairs up to 160-170 µm long. Abdominal plate with
2-3 pairs of long stout submarginal hairs on anterior half, but with the
most posterior pair also large (Figs 58E, F) . . . . . . . . . . . . . . . . . . . . . . 5
5 Aptera with thick dorsal prosomal hairs having acuminate apices (Fig.
58A). No small fine dorsal prosomal hairs. Abdominal plate with only 2
long stout pairs of submarginal hairs on anterior half (Fig. 58E) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneuraphis gallarum
- Aptera with thick dorsal prosomal hairs having fine-pointed apices
(Fig. 58B), and numerous small fine hairs (40-60 µm long) also present.
Abdominal plate with first 3 pairs of submarginal hairs all long and stout
(Fig. 58F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneuraphis litseicola
232 Litsea

6 Aptera with dorsal hairs of prosoma less than 2 x middle diameter of

ANT III, and blunt or rounded at apices (Fig. 58C) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis litseae
- Aptera with dorsal hairs of prosoma long and fine, 3-4 x middle diameter
of ANT III, and pointed at apices . . . . . . . . . . . . . Nipponaphis manoji
7 SIPH as large pores on shallow cones, surrounded by long hairs . . . 8
- SIPH tubular, much longer than basal width . . . . . . . . . . . . . . . . . . . 10
8 Aptera with SIPH cones pale and dorsal abdomen without pigmented
sclerites. R IV+V 0.8-0.9 x HT II . . . . . . . . . . . . . Aiceona robustiseta
- Aptera with SIPH cones dark and with small dark, hair-bearing sclerites
on dorsal abdomen. R IV+V 1.0-1.25 x HT II . . . . . . . . . . . . . . . . . . 9
9 ANT PT/BASE 0.25-0.45. ABD TERG 7 with 11-22 hairs, and ABD
TERG 8 with 8-14 hairs . . . . . . . . . . . . . . . . . . . . . . Aiceona titabarensis
- ANT PT/BASE 0.48-0.65. ABD TERG 7 with 6-10 hairs, ABD TERG
8 with 4-9 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona actinodaphnis
10 SIPH without hairs and similar in length to the large dark pointed CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinomegoura citricola
- SIPH much longer than CAUDA with numerous long hairs, CAUDA
helmet-shaped or broadly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 Antennae 4- or 5-segmented. First tarsal segments with 5 hairs. SIPH of
aptera tapering, broader at base than in middle . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Greenideoida (Pentatrichosiphum) lutea
- Antennae 5- or 6-segmented. First tarsal segments with 7 hairs. SIPH
of aptera cigar-shaped, broader in middle than at base . . . . . . . . . . 12
12 Antennae 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Antennae 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13 Abdominal tergum spinulose. R IV+V 2.1-2.7 x HT II . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum sankari
- Abdominal tergum smooth. R IV+V 1.4-1.8 x HT II . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum makii
14 ANT PT/BASE 1.6-1.9. SIPH dark, with interspersed long and short
hairs over entire length (e.g. Fig. 92G) . . . . . . Eutrichosiphum dubium
- ANT PT/BASE about 0.8. SIPH pale, with shorter hairs only near base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum litseae*
 Macadamia 233

LIVISTONA Palmae

Livistona altissima (?) Astegopteryx rhapidis

L. chinensis Cerataphis ?variabilis
(For palm aphids see B & E, 1984)

LONCHOCARPUS Leguminosae

Aphis craccivora

LOXOSTYLIS Anacardiaceae

Loxostylis alata Macrosiphum euphorbiae

MAACKIA Leguminosae

Maackia chinensis Myzus persicae

MACADAMIA Proteaceae

Aphis gossypii
Myzus persicae
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)
234 Macaranga

MACARANGA Euphorbiaceae

Macaranga tanaria Toxoptera aurantii

MACHILUS = PERSEA

MACLURA Moraceae

Maclura cochindriensis Toxoptera citricidus

M. javanensis Toxoptera citricidus
M. pomifera Aphis fabae
Myzus persicae
(Use key to polyphagous aphids, p. 532)

MAGNOLIA Magnoliaceae

Host Plant List

Magnolia denudata Aulacorthum magnoliae
M. fuscata [Nipponaphis karatanei]
M. kobus Neocalaphis magnoliae
M. liliflora Aulacorthum magnoliae
Neocalaphis magnoliae
Toxoptera odinae
M. obovata Neocalaphis magnolicolens
M. sieboldii Aulacorthum solani
Magnolia spp. Aulacorthum (Neomyzus)
circumflexum
Formosaphis micheliae
Macrosiphum euphorbiae
Myzus persicae
Tinocallis magnoliae
 Malacantha 235

Fig. 59. ANT III of A, Neocalaphis magnoliae and B, N. magnolicolens.

Key (o aphids on Magnolia

1 SIPH absent. Antennae of both aptera and alata 5-segmented. ANT I I I ,
IV and V of alata each with a single complex rhinarial structure occupying
most of the segment (Fig. 61 A) . . . . . . . . . . . . . . Formosaphis micheliae
SIPH present as tubes or small truncate cones . . . . . . . . . . . . . . . . . . . 2
2 SIPH tubular, longer than their basal widths, anal plate entire and
CAUDA tongue-shaped. (Viviparae apterous or alate) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
SIPH as small truncate cones, not longer than their basal widths. CAUDA
knobbed, anal plate bilobed. (All viviparae alate) . . . . . . . . . . . . . . . . 3
3 ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . Tinocallis magnoliae
- ANT PT/BASE more than 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 ANT III with two broad black bands, one in the middle and one distally
(Fig. 59A). All tibiae with black bases. Forewing veins heavily bordered
with fuscous, especially distally . . . . . . . . . . . . . Neocalaphis magnoliae
ANT III only gradually darkening distally (Fig. 59B). Tibiae without dark
basal sections, and forewing veins not bordered with fuscous . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neocalaphis magnolicolens

MALACANTHA = POUTERIA
236 Mallotus

MALLOTUS Euphorbiaceae

Mallotus japonicus Myzus persicae

Stomaphis malloti
M. oppositifolius Toxoplera aurantii

Rostrum longer than body. BL of adult more than 5 mm . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis malloti
Rostrum much shorter than body. BL of adult less than 4 mm . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

MALPIGHIA Malpighiaceae

Aphis craccivora, gossypii,

spiraecola
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

MALUS Apples, Crab-apples Rosaceae

A key to apple aphids is provided in B & E, 1984. Here we provide a more

comprehensive treatment of the aphid fauna of the genus Malus. The separa-
tion of Dysaphis species applies only to spring forms, and is based largely on
the work of Stroyan (1963) and Shaposhnikov (1986); the difficult taxonomy
of this group is discussed on p. 666.

Host Plant List

Malus angustifolia Aphis spiraecola
(Southern Crab-apple) [Hyalomyzus eriobotryae]
Muscaphis smithi
Prociphilus caryae ssp. fitchii
M. baccata Dysaphis sp. near anthrisci
(Siberian Crab-apple) Nearctaphis bakeri
Ovatus crataegarius, malisuctus
Prociphilus kuwanai
 Malus 237

M. coronaria Aphis gossypii, pomi

(Sweet Crab-apple)
M. floribunda Dysaphis devecta
Rhopalosiphum insertum
M. fusca Eriosoma lanigerum
(Oregon Crab-apple)
M. ionensis Aphis pomi
(Prairie Crab-apple) Dysaphis plantaginea
Rhopalosiphum insertum
M. orientalis Dysaphis affinis, anthrisci
majkopica, armeniaca,
brachycyclica, brancoi,
chaerophyllina, plantaginea,
radicola
M. pallasiana Dysaphis mordvilkoi, sibirica
M. pumila Allocotaphis quaestionis
(Apple) Aphidounguis mali
Aphis craccivora, eugeniae, fabae,
gossypii, pomi, spiraecola
Dysaphis anthrisci, a. majkopica,
armeniaca, brachycyclica,
brancoi, brancoi ssp. rogersoni,
bunii, chaerophylii,
chaerophyllina, devecta, flava,
malidauci, meridialis, mordvilkoi,
physocaulis, plantaginea,
radicola, sibirica, zini
Eriosoma lanigerum
[Hyalomyzus eriobotryae]
Longistigma xizangensis
Macrosiphum euphorbiae, rosae
Myzus persicae
Nearctaphis bakeri
Ovatus crataegarius, insitus,
malisuctus
Prociphilus caryae ssp. fitchii,
[crataegicola], kuwanai, oriens,
sasakii
Pterochloroides persicae
Pyrolachnus pyri
[Radisectaphis gyirongensis]
Rhopalosiphum insertum
Schizoneurella indica
Toxoptera aurantii, odinae
238 Malus

M. sieboldii Ovatus malisuctus

(Toringo Crab-apple)
M. sieversii Dysaphis affinis
M. silvestris = M. pumila
Malus sp. Dysaphis brachycyclica

Key to aphids on Malus

1 ANT PT/BASE less than 0.5. CAUDA broadly rounded, usually shorter
than width at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE more than 0.5 (usually more than 1). CAUDA semi-
circular, helmet-shaped or tongue-shaped, about as long as its basal width
or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2 Aptera, and sometimes alata also, with conspicuous wax pore plates.
SIPH either absent or present only as pores . . . . . . . . . . . . . . . . . . . . . 3
- Wax pore plates absent. SIPH in form of large, broad, dark, hairy cones
............................................................. 9
3 Wax pore plates of aptera comprising rings of circular, polygonal or
elongate facets surrounding single or multiple clear central areas. Wax
glands of alata small and indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Wax pore plates honeycomb-like, without any clear central areas. Alata
with well-developed wax pore plates, at least on thorax, or head and
thorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4 Antenna of aptera 4-segmented (Fig. 60A). Tarsi of aptera with only one
claw developed. Alata with unbranched media in forewing . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphidounguis mali
- Antenna of aptera 5- or 6-segmented, and tarsi with both claws equally
developed. Forewing of alata with once-branched media . . . . . . . . . . 5
5 Aptera without SIPH. Antenna of aptera 5-segmented, less than
0.17 x BL, with last 2 segments each about as broad as long. Wax pore
plates comprising groups of elongate facets radiating from 2-9 small clear
circular central areas (Fig. 60B) . . . . . . . . . . . . . . . Schizoneurella indica
- Aptera with SIPH as slightly raised pores with partly sclerotized rims.
Antenna of aptera usually 6-segmented, more than 0.17 x BL, with last
2 segments much longer than broad. Wax pore plates each comprising
large circular or polygonal facets enclosing one or more clear central areas
(Fig. 60C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanigerum
6 BL of spring migrant alata from leaf-nest gall (s.m.a.) 4mm or more.
R IV+V more than 0.2mm long and bearing 7 or more accessory hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
 Malus 239

Fig. 60. A, antenna of Aphidounguis mali (aptera); dorsal abdominal wax pore plate of B, Schizoneurella
indica and C, Eriosoma lanigerum; head of spring migrant alata of D, Prociphilus caryae fitchii and
E, P. sasakii; SIPH of apterae of F, Ovatus malisuctus, G, 0. crataegarius and H, Rhopalosiphum
insertum; hairs on HT I of I, Aphis eugeniae and J, A. pomi.

- BL of s.m.a. less than 4 mm. R IV+V less than 0.16 mm long and bearing
2-5 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 S.m.a. without any clear wax pore plates on head, and secondary rhinaria
on ANT V numbering 8-10 . . . . . . . . . . . . . . . . . . . Prociphilus kuwanai
- S.m.a. with 1-2 pairs of very pale and clearly-defined wax pore plates on
head; secondary rhinaria on ANT V numbering 0-7 . . . . . . . . . . . . . . 8
8 S.m.a. with one pair of circular wax pore plates on vertex of head. ANT
I (outside length) as long as or longer than its basal width (Fig. 60E). SIPH
present as small pigmented rings . . . . . . . . . . . . . . . . Prociphilus sasakii
240 Malus

- S.m.a. with 2 pairs of cephalic wax pore plates, one pair on frons and
the other on vertex, and ANT I shorter than its basal width (Fig. 60D).
SIPH absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus caryae fitchii
9 Dorsal abdomen with paired, pigmented spinal tubercles on each segment.
Alata with maculate forewings and short pterostigma . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterochloroides persicae
- Dorsal abdomen without dark tubercles. Forewings of alata not maculate,
with a long pterostigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Forewing of alata with pterostigma curved distally, extended around wing-
tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma xizangensis*
- Forewing of alata with pterostigma almost straight, not extending around
tip of wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrolachnus pyri
11 Head with well-developed antennal tubercles, their inner faces scabrous
or spinulose and bearing short, blunt or slightly clavate hairs (e.g.
Fig.42I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- Head with antennal tubercles either undeveloped or with inner faces
smooth and divergent and bearing pointed hairs . . . . . . . . . . . . . . . . . 15
12 SIPH moderately swollen on distal half (Fig. 121H) . . . . Myzus persicae
- SIPH cylindrical or tapering on distal half, or at most slightly swollen
subapically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Genital plate enlarged and produced posteriorly. ANT PT/BASE less than
4. SIPH dark, very coarsely imbricated (scabrous), and with a small flange
(Fig. 60F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus malisuctus
- Genital plate normal. ANT PT/BASE more than 4. SIPH pale or dusky,
moderately imbricated, with a well-developed flange (Fig. 60G) . . . . . 14
14 Spring migrant alata with 60-83 secondary rhinaria on ANT III, 36-52
on ANT IV and 13-22 on ANT V . . . . . . . . . . . . . . . . . . Ovatus insitus
- Spring migrant alata with 22-49 rhinaria on ANT III, 5-20 on IV and 0-10
on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus crataegarius
15 Head capsule of aptera spinulose or nodulose . . . . . . . . . . . . . . . . . . . 16
- Head capsule of aptera smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 SIPH short, of similar length to short CAUDA, with closely-spaced rows
of nodules or spinules (Fig. 42M) . . . . . . . . . . . . . . . Nearctaphis bakeri
- SIPH more than 4 x CAUDA; jet black and imbricated in aptera
(= fundatrix), dark and mainly smooth in alata . . . . Muscaphis smithi
17 CAUDA tongue- or finger-shaped, clearly longer than basal width . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
- CAUDA short, helmet-shaped, semicircular or triangular, not longer than
or about as long as its basal width in dorsal view . . . . . . . . . . . . . . . 27
 Malus 241

18 Body oval, with SIPH only about half as long as the distance between
their bases, without polygonal reticulation. Antennal tubercles weakly
developed or undeveloped. Aptera without rhinaria on ANT III . . . . 19
- Body spindle-shaped, SIPH longer than the distance between their bases,
with a subapical zone of polygonal reticulation. Antennal tubercles diver-
gent in dorsal view. Aptera with a few rhinaria on ANT III near base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
19 SIPH slightly swollen subapically and constricted before the well-
developed apical flange (Fig. 60H). Marginal tubercles on ABD TERG 7
placed posteriodorsally to spiracle, and no larger than the spiracular open-
ing (Fig. 42H) . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum insertum
- SIPH tapering from base to apex, with flange only moderately developed.
Marginal tubercles on ABD TERG 7 placed posterioventrally to spiracle
and usually larger than spiracular opening (Fig. 42H) . . . . . . . . . . . . 20
20 Stridulatory apparatus present, with sclerotic ridges on abdominal ster-
nites 5 and 6 and a row of peg-like hairs on hind tibia (Fig. 122). ANT
PT/BASE 3.5-5.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- No Stridulatory ridges present and peg-like hairs on hind tibia only in one
species (Aphis eugeniae). ANT PT/BASE less than 3.5 . . . . . . . . . . 21
21 Femoral hairs long and fine, many exceeding the length of the trochantero-
femoral suture. CAUDA with 6-24 hairs . . . . . . . . . . . . . . . . . . . . . . . 22
- Femoral hairs mostly short, with only a few ventral hairs long and those
not exceeding the length of the trochantero-femoral suture. CAUDA with
4-8 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
22 Aptera with black transverse bands on ABD TERG 7 and 8, and at least
some dark spinopleural markings anterior to SIPH. Alata with a series
of dorsal abdominal transverse bars. CAUDA tongue-shaped, without a
trace of a constriction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis fabae
- Aptera without dark dorsal abdominal markings, and alata with only
marginal sclerites. CAUDA finger-shaped, with a slight constriction near
midpoint (e.g. Fig.121T) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 Hind tarsus with 3 hairs (a sense peg plus 2 lateral hairs), like the fore
and mid-tarsi (Fig. 60I). Basal half of hind tibia with a spaced-out row
of short peg-like hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis eugeniae
- Hind tarsus with only 2 hairs (no sense peg; Fig. 60J). Hind tibia without
peg-like hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Marginal tubercles present on ABD TERG 2-4. CAUDA with 10-19 hairs
(rarely less than 13). R IV+V more than 120 µm . . . . . . . . Aphis pomi
- Marginal tubercles usually absent from ABD TERG 2-4. CAUDA with
7-15 hairs (rarely more than 12). R IV+V less than 120 µm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
242 Malus

25 Dorsum of aptera mainly black. SIPH and CAUDA black, the CAUDA
having a rather pointed apex . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
- Dorsum of aptera pale. SIPH usually darker than CAUDA, which has a
rounded apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
26 Front of head pale or dusky. SIPH of aptera pale at least on basal half,
sometimes dusky towards apex . . . . . . . . . . . . Macrosiphum euphorbiae
- Front of head black. SIPH wholly black . . . . . . . . Macrosiphum rosae
27 Abdomen without marginal tubercles. SIPH more than 3.5 x CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allocotaphis quaestionis
- Abdomen with marginal tubercles on most segments. SIPH less than
3.5 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 Antennae of aptera at least as long as distance from frons to bases of
SIPH, and those of alata about as long as body. Aptera without pigmenta-
tion of abdominal tergites anterior to SIPH . . . . . Dysaphis plantaginea
- Antennae of aptera shorter than the distance from frons to bases of SIPH,
and those of alata less than body length. Aptera with dorsal abdominal
pigment spots or patches anterior to SIPH . . . . . . . . . . . . . . . . . . . . . 29
29 Colonies include many apterae (or alatiform apterae with sclerotized
thorax) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
- Colonies comprise mainly immature and adult alatae . . . . . . . . . . . . 31
30 Longest hairs on ANT III longer than basal diameter of segment. True
aptera (no sclerotization of thorax) with only sparse abdominal pigmenta-
tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis devecta
- Longest hairs on ANT III less than 0.5 x diameter of base of segment.
True aptera with broad dark bands or paired patches on all tergites . . . .
................................................ Dysaphis affinis
31 (Remaining couplets apply only to spring migrant alatae) Each antenna
with a total of 112-198 secondary rhinaria. Marginal tubercles usually
present on ABD TERG 1-7 inclusive, sometimes absent from ABD TERG
6. HT I with 3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis brancoi
- Each antenna with a total of 20-113 secondary rhinaria. Marginal
tubercles present on ABD TERG 1-5, 1-5 and 7, or 1-7. HT I with 2 or
3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 Ratio of total number of secondary rhinaria on both antennae to length
of SIPH (in µm) 1.12-1.64 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
- Ratio of total number of secondary rhinaria on both antennae to length
of SIPH (in µm) 0.26-1.00 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33 Marginal tubercles frequently present on ABD TERG 1-7 inclusive.
Ratios of diameter of SIPH at midpoint to diameters of largest and
 Malus 243

smallest marginal tubercles on ABD TERG 1-5 are 0.8-1.1 and 1.1-1.8,
respectively. ANT V with 0-6 secondary rhinaria . . . . . Dysaphis sibirica
- Marginal tubercles present on ABD TERG 1-5, or 1-5 and 7 (never on
6). Ratios of diameter of SIPH at midpoint to diameters of largest and
smallest tubercles on ABD TERG 1-5 are 1.3-2.1 and 2.7-6.0, respec-
tively. ANT V with 3-10 secondary rhinaria . . . . Dysaphis mordvilkoi
34 Posterior part of pronotum with a pair of pleural hairs . . . . . . . . . . 35
- Posterior part of pronotum without a pair of pleural hairs (although occa-
sionally one is present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
35 ABD TERG 8 with 7-10 hairs. Longest hair on ANT III 42-64 µm long,
1.6-2.7 x basal diameter of segment . . . . . . . . . . Dysaphis chaerophylli
- ABD TERG 8 with 4-5 hairs. Hairs on ANT III maximally 39µm, not
more than 1.7 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . 36
36 CAUDA with 6-9 hairs. Longest hair on ANT III is 13-18 µm, usually
shorter than basal diameter of segment . . . . . . . . . . . . . . Dysaphis flava
- CAUDA with 4-6 hairs, usually 5; rarely with 7. Longest hair on ANT
I I I is 9-39µm, longer or shorter than basal diameter of segment, but if
shorter then very b l u n t or dilated apically . . . . . . . . . . . . . . . . . . . . . . 37
37 Spinopleural hairs on ABD TERG 3 usually short and blunt, maximally
ll-23µm long. Longest hair on ANT III 9-29 µm, 0.4-1.3 x basal
diameter of segment. Marginal tubercles present on ABD TERG 1-7
inclusive (occasionally absent from one side on 6) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis radicola group
- Spinopleural hairs on ABD TERG 3 pointed, the longest being more than
25 µm long. Longest hair on ANT III in range 20-39 µm, 1.0-1.7 x basal
diameter of segment. Marginal tubercles present on ABD TERG 1-5, or
1-5 and 7; absent from 6 except rarely on one side only . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis anthrisci group
38 Each antenna with a total of 60-99 secondary rhinaria. R IV+V less than
1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis affinis
- Each antenna with a total of 26-59 secondary rhinaria. R IV+V usually
more than 1.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39 Total number of 'spinal' tubercles (including those on head) 6-19. Each
antenna with a total of 37-59 (usually more than 46) secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis armeniaca
- Total number of 'spinal' tubercles (including those on head) 2-7. Each
antenna with a total of 26-46 secondary rhinaria . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis foeniculus malidauci
244 Mammea

MAMMEA Guttiferae

Mammea americana Aphis gossypii

Toxoptera aurantii
(See key to polyphagous aphids, p, 532)

MANGIFERA Mangoes Anacardiaceae

Aphis craccivora, fabae/solanella,

gossypii, spiraecola
Greenidea mangiferae
Macrosiphum centranthi
Sinomegoura citricola
Sitobion krahi
Toxoptera aurantii, odinae

Key to aphids on Mangifera

1 SIPH with numerous very long hairs . . . . . . . . . Greenidea mangiferae
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH with a subapical zone of polygonal reticulation . . . . . . . . . . . . . 3
SIPH without any polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
3 SIPH and antennae mainly dark in both aptera and alata. SIPH at mid-
point of similar thickness to hind femora. Hairs on ANT III all less than
half basal diameter of segment . . . . . . . . . . . . . . . . . . . . . Sitobion krahi
SIPH and antennae mainly pale in aptera, dusky to dark in alata. SIPH
at midpoint much thinner than hind femora. Hairs on ANT III maximally
more than half basal diameter of segment . . . . Macrosiphum centranthi

MARGARETTA Asclepiadaceae

Margaretta whitei Aphis nerii

 Melaleuca 245

MARKHAMIA Bignoniaceae

Markhamia acuminata Aphis craccivora, gossypii

M. platycalyx Myzus persicae
(Use key to polyphagous aphids, p. 532)

MAYTENUS Celastraceae

Macrosiphum euphorbiae
Myzus persicae
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

MELALEUCA Myrtaceae

Melaleuca leucodendron Aphis gossypii

Greenidea (Trichosiphum)
formosana
Myzus persicae
M. teretifolia Taiwanaphis sp.
Melaleuca sp(p). Myzus persicae
Toxoptera aurantii

Key to aphids on Melaleuca

1 SIPH with numerous long hairs. Dorsum of aptera dark, sclerotic, bearing
long hairs with furcate apices. Hind tibiae with imbrications bearing
minute spinules . . . . . . . . . . . . . . . Greenidea (Trichosiphum) formosana
SIPH without hairs. Dorsum pale and membranous bearing hairs with
finely pointed or blunt apices. Hind tibiae without spinulose imbrication
............................................................. 2
2 SIPH as small, broad-based cones. CAUDA knobbed, anal plate bilobed.
ABD TERG 7 and 8 of aptera each with 2 pairs of backwardly-directed
processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis sp(p).
246 Melia

- SIPH tubular. CAUDA tongue- or finger-shaped. ABD TERG 7 and 8

without processes . . . . . . . . . . go to key to polyphagous aphids, p. 532

MELIA Meliaceae

Aphis gossypii, spiraecola

Myzus ornatus
[Sinomegoura nepalensis Das and
Raychaudhuri, 1983]
(Use key to polyphagous aphids, p. 532)

MELIOSMA Meliosmaceae

Meliosma myriantha Mollitrichosiphum yamabiwae

M. rhoifolia Mollitrichosiphum taiwanum
M. rigida Mollitrichosiphum taiwanum,
yamabiwae
M. tennis Mollitrichosiphum yamabiwae

Key to aphids on Meliosma

The 2 aphid species, which are both in Mollitrichosiphum subgenus Meta-
trichosiphon, can be separated by the following couplet:
Aptera with dark ventral abdominal patch. Alata with less than 20
rhinaria on ANT III . . . . . . . . . . . . . . . . . Mollitrichosiphum yamabiwae
Aptera without dark ventral abdominal patch, Alata with 20 or more
rhinaria on ANT III . . . . . . . . . . . . . . . . . . Mollitrichosiphum taiwanum

MEMECYLON Melastomataceae

Brachycaudus helichrysi
Taiwanaphis memecyloni

Key to aphids on Memecylon

CAUDA knobbed, anal plate bilobed. Posterior abdominal tergites of 4th
 Metroxylon 247

instar nymph with backwardly-directed, curved processes (apterous

morph unknown) . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis memecyloni
CAUDA semicircular, without a constriction; anal plate entire. Abdomen
without processes . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus helichrysi
(or try key to polyphagous aphids, p. 532)

MESUA Guttiferae

Mesua ferrea Greenideoida ceyloniae

(Na, Ironwood) Toxoptera aurantii

Key to aphids on Mesua

Body elongate, more than twice as long as wide. SIPH more than
0.5 x BL with numerous long hairs . . . . . . . . . Greenideoida ceyloniae
Body oval. SIPH less than 0.2 x BL, without hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
(or try key to polyphagous aphids, p. 532)

METASEQUOIA Taxodiaceae

Metasequoia glyptostroboides Illinoia morrisoni

METROSIDEROS Myrtaceae

Metrosideros lucida Myzus persicae

METROXYLON Palmae

Metroxylon sagu Cerataphis variabilis

(For palm aphids see B & E, 1984)
248 Michelia

MICHELIA Magnoliaceae

Aphis spiraecola
Formosaphis micheliae
Myzus persicae
Prociphilus micheliae
Sinomegoura citricola

Key to aphids on Michelia

1 SIPH absent. ANT PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . 2
- SIPH present. ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
2 Antenna of alata 5-segmented; ANT III, IV and V each with an extensive,
complex rhinarial structure occupying most of segment (Fig. 61A) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Formosaphis micheliae
Antenna of alata 6-segmented, with separate, transversely elongate secon-
dary rhinaria on ANT III and IV only, fringed with minute spinules
(Fig. 61B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus micheliae

MICROCOS Tiliaceae

Microcos paniculata Cervaphis schoutedeniae

MILLETTIA Leguminosae

Millettia eetveldiana Pterasthenia matileae

M. reticulata Aulacophoroides formosanum
M. thonningii Aphis craccivora
Millettia sp. Acyrthosiphon pisum

Key to aphids on Millettia

1 All viviparae alate, with knobbed CAUDA and bilobed anal plate . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterasthenia matileae
 Mimosa 249

Fig. 61. Antennae of alatae of A, Formosaphis micheliae and B, Prociphilus micheliae.

Viviparae in colonies often mostly apterous; CAUDA tongue-like or

finger-like, anal plate entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Dorsal abdomen of aptera with a large black sclerotic patch. Frons with
antennal tubercles undeveloped. CAUDA dark like SIPH . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
Dorsal abdomen dusky or pale in cleared specimens. Antennal tubercles
well developed, divergent. CAUDA pale . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 SIPH black, as thick as CAUDA and swollen in middle, arising from large
dark sclerites. Head, antennae and distal parts of femora dark, all with
rather long hairs . . . . . . . . . . . . . . . . . . . . . . Aulacophoroides formosana
SIPH pale, attenuate, much thinner than CAUDA, with pale bases. Head,
antennae and femora all mainly pale with very short hairs . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon pisum

MIMOSA Leguminosae

Mimosa pudica Sitobion mimosae

(Sensitive Plant)
Mimosa sp. Aphis craccivora
250 Mimusops

Key to aphids on Mimosa

- SIPH brown and reticulated over distal 20%. CAUDA pale. ANT
PT/BASE 4.0-5.75 . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion mimosae*
SIPH black and without polygonal reticulation. CAUDA black. ANT
PT/BASE 1.9-2.8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
(or try key to polyphagous aphids, p. 532)

MIMUSOPS Sapotaceae

Mimusops zeyheri Toxoptera aurantii

MONOTES Dipterocarpaceae

Monotes caloneurus Aphis craccivora, gossypii

[Sitobion sp.]
M. loandensis Aphis gossypii
(Use key to polyphagous aphids, p. 532)

MORINDA Rubiaceae

Aphis gossypii

MORINGA Moringaceae

Moringa oleifera Aphis spiraecola

Myzus persicae
(Use key to polyphagous aphids, p. 532)
 Myrica 251

MORUS Mulberry Moraceae

Aphis craccivora, gossypii,

spiraecola
[Aphis mori Clarke, 1903]
Toxoptera aurantii, citricidus
(Use key to polyphagous aphids, p. 532)

MUNDULEA Leguminosae

Mundulea sericea Aphis craccivora

MURRAYA Rutaceae

Aphis spiraecola
Myzus persicae
Sinomegoura citricola
Toxoptera aurantii, citricidus
(Use key to polyphagous aphids, p. 532)

MYRICA Myrtles Myricaceae

Host Plant List

Myrica asplenifolia Calaphis myricae
Greenidea (Trichosiphum) myricae
Illinoia azaleae, borealis
M. californica Aphis fabae
M. cerifera Longistigma caryae
M. gale Calaphis myricae
Greenidea (Trichosiphum) myricae
Illinoia canadensis
252 Myrica

Macrosiphum euphorbiae
Myzocallis myricae
M. rubra Aulacorthum solani
Parathoracaphis cheni

The small but very diverse aphid fauna of Myrica consists mainly of species
recorded from shrubs such as M. gale, but these are nevertheless included in
the following key for the sake of completeness.

Key to aphids on Myrica

1 Aptera aleyrodiform; black, strongly sclerotized, with legs and antennae
concealed beneath body . . . . . . . . . . . . . . . . . . . . . Parathoracaphis cheni
Aptera of normal aphid form, with well-developed legs and antennae
............................................................. 2
2 BL more than 4.5mm. SIPH as large pores on broad dark hairy cones.
ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . . Longistigma caryae
BL less than 3.5mm. SIPH tubular or conical. ANT PT/BASE more
than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 SIPH tubular, with numerous long hairs . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) myricae
SIPH tubular or conical, w i t h o u t hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 SIPH in form of short truncated cones. CAUDA with a constriction, and
anal plate bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
SIPH tubular, tapering, cylindrical or swollen on distal half. CAUDA
finger-like, without a constriction, and anal plate entire . . . . . . . . . . . 6
5 Antennae shorter than body, with ANT PT/BASE less than 2. Head and
thorax with longitudinal black stripes, and abdomen with paired dark
segmental patches. Tibiae with similar pigmentation to distal parts of
femora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis myricae
Antennae longer than body, with ANT PT/BASE more than 2. Dorsum
without a pattern of dark markings. Tibiae black, contrasting with paler
femora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calaphis myricae
6 SIPH tapering or cylindrical on distal half (or, if clearly swollen, then
without subapical reticulation) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
SIPH clearly swollen on distal half, with polygonal reticulation distal to
swelling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 First tarsal segments with 5 hairs . . . . . . . . . . . . . . . . . . . Illinoia azaleae
First tarsal segments with 3 hairs (occasionally with 4, but never 5). . . .
............................................................. 8
 Nectandra 253

8 SIPH dark over most or whole of length, with a small flange. HT II only
0.07-0.10 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia borealis
SIPH pale or dusky, or only dark at apices, with a well-developed flange.
HT II 0.10-0.14 mm long . . . . . . . . . . . . . . . . . . . . . . Illinoia canadensis

MYRTUS Myrtaceae

Aulacorthum (Neomyzus)
circumflexum
Myzus persicae
Toxoptera aurantii
(See key to polyphagous aphids, p. 532)

NAUCLEA Rubiaceae

Nauclea esculenta Aphis gossypii

NECTANDRA Lauraceae

Nectandra antillana Toxoptera aurantii

Nectandra sp. Lizerius tuberculatus

- ANT PT/BASE less than 1. Anal plate bilobed, CAUDA with an apical
knob. Body of apterae with numerous club-shaped tubercular processes
(Fig. 63B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lizerius tuberculatus
ANT PT/BASE more than 3. Anal plate entire, CAUDA tongue-shaped.
Body of aptera without club-shaped processes . . . . . Toxoptera aurantii
(or try key to polyphagous aphids, p. 532)
254 Neolitsea

NEOLITSEA Lauraceae

Neolitsea sericea Machilaphis machili

Nipponaphis coreanus, litseae

Although only these three species are recorded from Neolitsea, it seems pro-
bable that it may be colonized by other aphids which occur on Machilus and
Litsea. Lists and keys for those plant genera should therefore be consulted.

NOTHOFAGUS Southern Beeches Fagaceae

Three related but distinct genera of Drepanosiphinae are found on Nothofagus:

Neuquenaphis and Neosensoriaphis (9 species) in South America and Taiwan-
aphis (subgenus Sensoriaphis; 4 species) in Australasia. Neuquenaphis similis
is only known from the apterous morph, and N. michelbacheri only from
the alate.

Host Plant List

Nothofagus alpina Neuquenaphis edwardsi,
michelbacheri, sensoriata
N. antarctica Neuquenaphis edwardsi,
michelbacheri
N. carrii Taiwanaphis niuginii
N, cunninghami Taiwanaphis tasmaniae
N. dombeyi Neuquenaphis bulbicauda,
chilensis, edwardsi,
michelbacheri, palliceps,
sensoriata
N. fusca Taiwanaphis nothofagi
N. glauca Neuquenaphis michelbacheri,
sensoriata
N. moorei Taiwanaphis furcifera
N. obliqua Neosensoriaphis parva
Neuquenaphis edwardsi, essigi,
palliceps, schlingeri, sensoriata,
similis
 Noíhofagus 255

N. pumilio Neuquenaphis edwardsi, similis

N. solandri var. cliffortiodes Taiwanaphis nothofagi
N. truncata Taiwanaphis nothofagi

Key to aphids on Nothofagus

1 ANT PT/BASE clearly less than l. Aptera with 2-8 backwardly directed
finger-like processes on ABD TERG 6-8; such processes absent or much
reduced on head, thorax and more anterior tergites, or in single marginal
pairs only on ABD TERG 1-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE equal to or greater than 1. Aptera either with long tapered
finger-like processes on head and at least marginally on all abdominal
tergites, or with such processes on ABD TERG 7 and 8 only . . . . . . 5
2 Aptera (fundatrix) with spinal processes on ABD TERG 8 fused at base
to form a forked structure (Fig. 62A). Alata with ANT III bearing 35-57
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis furcifera
- Aptera with spinal processes on ABD TERG 8 separate. Alata with ANT
I I I bearing 9-20 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ANT PT/BASE more than 0.5. Aptera with only 1 pair of large (spinal)
processes on ABD TERG 8, marginal processes being undeveloped
(Fig. 62B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis niuginii
- ANT PT/BASE less than 0.5. Aptera with marginal processes at least on
ABD TERG 6 and 7, as well as the spinal processes on ABD TERG 8
............................................................. 4
4 Aptera with marginal as well as spinal processes on ABD TERG 8
(Fig. 62C), and also with finger-like marginal tubercles on ABD TERG
1-4. Body broadly oval. Dorsal abdomen (aptera and alata) with small
dark markings but mainly unsclerotized . . . . . . Taiwanaphis tasmaniae
- Aptera with marginal processes only well developed on ABD TERG 7
(Fig. 62D), or 6 and 7. Body narrowly oval, aptera and alata both with
transverse dark sclerotic bars . . . . . . . . . . . . . . . Taiwanaphis nothofagi
5 Head of aptera without any finger-like processes. ABD TERG 7 with 2
pairs of finger-like processes, and ABD TERG 8 with 1 pair (Fig. 62E).
Alata without finger-like abdominal processes, and with ANT PT/BASE
less than 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosensoriaphis parva
- Head of aptera with 1-2 pairs of finger-like processes, and ABD TERG
1-8 all with at least marginal processes, if alata is entirely lacking finger-
like abdominal spinal processes then ANT PT/BASE is more than 4
............................................................. 6
6 Head of aptera with 1 pair of long, finger-like processes (Fig. 62G);
abdomen with spinal processes much reduced or absent, and marginal
256 Northofagus

Fig. 62. Processes on ABD TERG 7 and 8 of aptera of A, Taiwanaphis furcifera (fundatrix),
B, T. niuginii, C, T. tasmaniae, D, T. nothofagi, E, Neosensoriaphis parva and F, Neuquenaphis chilensis',
processes on head of aptera of G, Neuquenaphis schlingen and H, Neuquenaphis essigi; processes on
ABD TERG 3 of aptera of I, Neuquenaphis essigi and J, N. sensoriata.
 Nothofagus 257

processes on each segment comprising 1 long pair, plus 1 (submarginal)

pair 0.3-0.5 as long. Alata with all abdominal spinal processes much
reduced (not longer than their basal widths), or absent . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis schlingen
- Head of aptera with at least 2 pairs of long processes; abdomen with spinal
as well as marginal processes long and finger-like. Alata with thin, finger-
like spinal processes at least on ABD TERG 1-3 . . . . . . . . . . . . . . . . . 7
7 ANT PT/BASE usually less than 1.5 in aptera, less than 2 in alata.
CAUDA with the knob either almost spherical, or rounded with a conical
apex (heart-shaped). Aptera with 2 pairs of finger-like processes on vertex
between eyes (e.g. Fig. 62H), as well as 1 or 2 more anterior pairs, and
ABD TERD 7 with 6-8 processes (e.g. Fig. 62F) . . . . . . . . . . . . . . . . . 8
- ANT PT/BASE usually greater than 1.5 in aptera, greater than 2 in alata.
CAUDA with an ellipsoid or tapering knob, clearly longer than its max-
imum width. Aptera with or without 1 pair of finger-like processes on
vertex between eyes (as well as 1 pair between antennal bases), and ABD
TERG 7 usually with 4 processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 CAUDA of aptera with knob a l i t t l e broader than long, with a broadly
rounded apex. Dorsal processes of aptera arising from rounded dark
sclerites (Fig. 62I) that coalesce into transverse bars on ABD TERG 6 and
7. Alata with forewing veins unbordered . . . . . . . . Neuquenaphis essigi
- CAUDA of aptera rounded basally but with an abruptly conical apex,
making it cordate or heart-shaped in dorsal view and a little longer than
its maximum width (Fig. 62F). Dorsal processes of aptera not arising from
round sclerites. Alata with forewing veins bordered with fuscous . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis chilensis
9 R IV+V very short, shorter than or about as long as its basal width.
CAUDA paler than SIPH . . . . . . . . . . . . . . . . . . Neuquenaphis palliceps
- R IV+V at least 1.3 x its basal width. CAUDA with similar pigmentation
to SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Apterae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Alatae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
11 R IV+V with 8-12 accessory hairs. ANT PT/BASE 1.5-1.9. Dorsal
abdominal processes arising from pigmented irregular sclerotic bars and
patches (Fig. 62J) . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis sensoriata
- R IV+V with 3-6 accessory hairs. ANT PT/BASE 2.4-4.5. Dorsal
abdomen pale, or dusky or very sclerotic, but without clearly defined
pigmented sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Antennae about 1.5 x BL. Embryos with capitate dorsal hairs 9-13 µm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis similis
258 Nyssa

- Antennae 0.6-1.1 x BL. Embryos with dorsal hairs less than 4µm long
(shorter than spinules on processes) . . . . . . . . . Neuquenaphis edwardsi
13 ANT PT/BASE 2.0-3.2. ANT III with 10-30 rhinaria. R IV+V with 7-12
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis sensoriata
- ANT PT/BASE 3.2-5.5, ANT III with 3-16 rhinaria. R IV+V with 4-8
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 ANT III with 8-16 rhinaria. Spinal processes on ABD TERG 1-3 pale and
mostly arising from unpigmented or only very weakly pigmented sclerites.
Forewing veins rather dark, the oblique veins beginning and ending in
conspicuous fuscous triangles (hand lens) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis michelbacheri
- ANT III with 3-9 rhinaria. Spinal processes on ABD TERG 1-3 pig-
mented, arising from pigmented sclerites. Forewing veins with very little
fuscous at ends . . . . . . . . . . . . . . . . . . . . . . . . . . . Neuquenaphis edwardsi

NYSSA Cornaceae

Nyssa sylvatica Aphis coreopsidis

Phylloxerina nyssae

OCHNA Ochnaceae

Sitobion ochnearum

OCHROMA Bombacaceae

Aphis gossypii

OCOTEA Lauraceae

Ocotea acutifolia Lizerius ocoteae

O. glomerata Lizerius tuberculatus
 Osmanthus 259

Fig. 63. End of abdomen (ABD TERG 6-8) of aptera of A, Lizerius ocoteae and B, L. tuberculatus.

Key to aphids on Ocotea

Antennae of aptera 5-segmented. Dorsal body membranous, with numer-
ous short spinulose club-shaped or crown-shaped processes. ABD TERG
8 with 4 finger-like processes (Fig. 63B). R IV+V longer than HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lizerius tuberculatus
Antenna of aptera 6-segmented. ABD TERG 1-5 each with 6 sclerites, the
marginal ones developed into short conical processes, and transverse bars
on ABD TERG 6-8; ABD TERG 8 bearing 2 large conical processes and
2-4 much smaller ones (Fig. 63A). R IV+V shorter than HT II . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lizerius ocoteae

OLEA Olives Oleaceae

Olea europaea Prociphilus oleae

[Tetraneura agnesii del Guercio,
1920]

OSMANTHUS Oleaceae

Osmanthus aquilifolium Prociphilus osmanthae

O. heterophyllus Prociphilus osmanthae
260 Ostrya

OSTRYA Hop-hornbeams Corylaceae

Ostrya carpinifolia Pterocallis ostryae

O. japonica Mesocallis pteleae
O. virginiana Macrosiphum (Neocorylobium)
pseudocoryli

Key to aphids on Ostrya

1 SIPH long, tubular, dark, with subapical zone of polygonal reticulation.
CAUDA long and tongue-shaped, anal plate entire. ANT PT/BASE more
than 5 . . . . . . . . . . . . . . . . Macrosiphum (Neocorylobium) pseudocoryli
SIPH as short, truncated cones. CAUDA knobbed, anal plate bilobed.
ANT PT/BASE 1 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT III of alata dark, with 7-14 secondary rhinaria. ANT PT/BASE
0.8-1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesocallis pteleae
ANT III of alata pale with 4-7 secondary rhinaria. ANT PT/BASE
0.3-0.6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocallis ostryae

OSTRYODERRIS = AGANOPE

OZOROA Anacardiaceae

Ozoroa insignis Aphis gossypii

O. paniculosa Aphis gossypii

PANAX Araliaceae

Toxoptera aurantii, odinae

(Use key to polyphagous aphids, p. 532)
 Paulownia 261

PANDANUS Pandanaceae

Pandanus tectorius Astegopteryx pandani

PAPPEA Sapindaceae

Pappea capensis Aphis gossypii

Toxoptera aurantii, citricidus
(Use key to polyphagous aphids, p. 532)

PARABENZOIN = UNDERA

PARINARIA Chrysobalanaceae

Aphis gossypii

PASANIA = LITHOCARPUS

PAULOWNIA Serophulariaceae

Aphis craccivora, gossypii,

spiraecola
Myzus persicae
(Use key to polyphagous aphids, p. 532)
262 Peltophorum

PELTOPHORUM Leguminosae

Peltophorum africanum Myzus persicae

P. fleragineum Tinocallis himalayensis

Key to aphids on Peltophorum

SIPH in form of short, truncate cones. CAUDA knobbed, anal plate
bilobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis himalayensis
SIPH tubular, slightly swollen on distal half. CAUDA tongue-shaped,
anal plate entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
(or try key to polyphagous aphids, p. 532)

PERSEA Lauraceae

Host Plant List

Persea americana Aphis gossypii, spiraecola
Myzus persicae
Sinomegoura citricola
Toxoptera aurantii
P. bombycana Schizoneuraphis himalayensis
P. bournei Longistigma xizangensis
Machilaphis machili
P. gamblei Aiceona retipennis
P. longipaniculata Machilaphis machili
P. odoratissima Aiceona pseudosugii
Machilaphis machili
P. thunbergi Machilaphis machili
(Pao-hua, Tabu, Tamagusa) Nipponaphis machili, machilicola
Schizoneuraphis machiliphaga
Persea sp(p). Nipponaphis amamiana
Sinonipponaphis formosana

Key to aphids on Persea

1 Aptera of normal aphid form, with well-developed legs and 6-segmented
antennae. Alata with circular secondary rhinaria . . . . . . . . . . . . . . . . . 2
 Persea 263

Aptera aleyrodiform with reduced legs, very short 3- or 4-segmented

antennae and variably fused head, thorax and anterior abdominal tergites.
Antenna of alata with annular secondary rhinaria . . . . . . . . . . . . . . . . 6
2 SIPH tubular, longer than basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
SIPH on flat hairy cones or merely as pores . . . . . . . . . . . . . . . . . . . . . 3
3 Body elongate oval. Hairs very sparse and mainly very short; those on
antenna much less than half basal diameter of ANT III. SIPH merely
small pores, not surrounded by hairs . . . . . . . . . . . Machilaphis machili
Body oval. Hairs long and dense; those on antennae longer than basal
diameter of ANT I I I . SIPH pores large, on hairy cones . . . . . . . . . . 4
4 BL more than 6mm. Eyes multifaceted. Forewing of alata with ptero-
stigma extending almost to wing-tip . . . . . . . . Longistigma xizangensis
BL less than 4 m m . Eyes of aptera 3-faceted. Forewing of alata with a
normal pterostigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 ANT PT/BASE 0.30-0.45. R IV+V 0.52-0.70 x HT II, and bearing 2-6
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona retipennis
- ANT PT/BASE 0.5-0.6. R IV+V about 0.8 x HT I I , and bearing 6-8
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona pseudosugii
6 Dorsal cuticle of aptera smooth, densely covered with short, thorn-like,
broad-based hair-like processes . . . . . . . . Sinonipponaphis formosana*
Dorsal cuticle of aptera either smooth or with densely pustulate ornamen-
tation; hairs of variable length, no thorn-like processes . . . . . . . . . . . 7
7 Spinal and submarginal hairs of aptera fine and not exceeding length of
antennae . . . . . . . . Nipponaphis spp. (amamiana, machili, machilicola)
Spinal and submarginal hairs of aptera mostly long and thick, the longest
exceeding the length of the antennae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Abdominal plate (fused ABD TERG 2-7) with first 3 pairs of submarginal
hairs very stout and dark, much longer and thicker than the posterior 3
pairs (Fig. 58D). Dorsal hairs of prosoma (i.e. fused head, thorax and
ABD TERG 1) often include some short, fine hairs as well as the paired,
stout spinal and submarginal hairs . . . . . Schizoneuraphis himalayensis
Abdominal plate with only the penultimate 2 pairs of submarginal hairs
(the fourth and fifth pairs) small, the last submarginal pair being almost
as large as the first 3 pairs (e.g. as in Fig. 58F) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneuraphis machiliphaga
264 Phellodendron

PHELLODENDRON Rutaceae

Phellodendron amurense Chaitophorus pheleodendroni

PHOEBE Lauraceae

Phoebe elongata Toxoptera aurantii

Ph. porphyria Lizerius ocoteae
Ph. sheareri Aiceona sp.
Machilaphis machili

Key to aphids on Phoebe

1 ANT PT/BASE 3.5-5.1. SIPH tubular. CAUDA tongue-shaped and
black like SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
ANT PT/BASE less than 0.5. SIPH as slightly raised pores or hairy cones.
CAUDA knobbed or broadly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 CAUDA with an elongate knob (Fig. 53A) . . . . . . . . . Lizerius ocoteae
CAUDA broadly rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Long-haired aphid, apterae with eyes of only 3 facets. SIPH on hairy cones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiceona sp.
Body hairs inconspicuous, apterae with compound eyes. SIPH as pores
ringed with a few hairs . . . . . . . . . . . . . . . . . . . . . . . Machilaphis machili

PHOTINIA Rosaceae

Host Plant List

Photinia arguta Nippolachnus bengalensis
Ph. X fraseri Brachycaudus helichrysi
Fimbriaphis gentneri
Macrosiphum euphorbiae
 Photinia 265

Ph. glabra Sinomegoura photiniae

Toxoptera aurantii

Ph. integrifolia Sinomegoura photiniae

Ph. notoniana Prociphilus sp.

Ph. Serrulata Aphis spiraecola .

(Chinese Hawthorn) Myzus persicae

Ph. variabilis Sinomegoura photiniae

Ph. villosa Aphis gossypii, spiraecola
Melanaphis bambusae
Prociphilus ushikoroshi

Photinia sp. [Greenidea photiniphaga

Raychaudhuri, Ghosh, Banerjee
and Ghosh, 1973]

Key to aphids on Photinia

1 ANT PT/BASE less than 0.5. CAUDA bluntly rounded . . . . . . . . . . 2

- ANT PT/BASE more t h a n 1. CAUDA tongue- or finger-like . . . . . . 3

2 Body and appendages with numerous long, fine hairs. SIPH pores large,
on hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . Nippolachnus bengalensis

Body and appendages with sparse, short hairs. SIPH pores indistinct or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus sp(p).

3 Antennal tubercles weakly developed. SIPH conical truncate or tapering

without trace of swelling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Antennal tubercles well developed, their inner faces divergent or con-

vergent in dorsal view. SIPH usually slightly swollen, especially on inner
sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

4 SIPH short and conical; in aptera less than twice as long as their basal
widths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

SIPH more than twice as long as their basal widths . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

5 CAUDA rounded, about as long as its basal width. R IV+V long, more
than twice as long as width at base . . . . . . . . . Brachycaudus helichrysi

CAUDA clearly longer than its width at base. R IV+V short; in alata
(emigrant) shorter that its basal width . . . . . . . . Melanaphis bambusae
266 Phyllanthus

6 SIPH dark, less than 1.5 x the long, pale, pointed CAUDA . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinomegoura photiniae
SIPH pale or dark only at tips, about twice length of CAUDA or more
............................................................. 7
7 SIPH with a subapical zone of polygonal reticulation. Alata without a
black dorsal abdominal patch . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH without any polygonal reticulation. Alata with a black dorsal
abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 ANT III of aptera with a few rhinaria near base. ANT PT/BASE ratio
5 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri
- ANT III of aptera without rhinaria. ANT PT/BASE usually less than 4
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae

PHYLLANTHUS Euphorbiaceae

Host Plant List

Phyllanthus discoideus Schoutedenia lutea
Ph. emblica Schoutedenia emblica
Sitobion takahashii
Ph. floribundus Aphis gossypii
Schoutedenia lutea
Ph. maderaspatensis Aphis gossypii
Sitobion ?mimosae, phyllanthi
Ph. muelleranus Schoutedenia lutea
Sitobion africana
Ph. myrtifolius Aphis eugeniae
Ph. reticulatus Schoutedenia lutea
Phyllanthus spp. Aphis craccivora
[Chuansicallis chengtuensis]
Eonaphis phyllanthi
Macrosiphum euphorbiae
Myzus ornatus, persicae
[Tinocallis nigropunctata]

Key to aphids on Phyllanthus

1 At least ABD TERG 7 with a pair of large, backwardly-pointed processes.
Antenna 5-segmented. Aptera with reduced eyes . . . . . . . . . . . . . . . . . . 2
 Phyllanthus 267

ABD TERG 7 without large processes. Antenna 6-segmented. Aptera with

well-developed multifaceted eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Head, thorax and ABD TERG 1-7 all with paired spinal processes, and
large marginal processes also present on at least ABD TERG 5 and 7. ANT
PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . Eonaphis phyllanthi
- Large processes on ABD TERG 7 only. ANT PT/BASE less than 1
.....................................:...................... 3
3 R IV+V shorter than PT. ANT PT/BASE more than 0.5 . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schoutedenia lutea
- R IV+V longer than PT. ANT PT/BASE usually less than 0.5 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schoutedenia emblica
4 SIPH with a subapical zone of polygonal reticulation (if this is very short,
then SIPH are constricted at apex with a very small flange) . . . . . . . 5
SIPH without any subapical polygonal reticulation . . . . . . . . . . . . . . . 9
5 Hairs on ANT I I I maximally more than 0.5 x diameter of ANT I I I at
midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
Hairs on ANT I I I very short, maximally much less than 0.5 x diameter
of ANT I I I at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 SIPH constricted at apex, w i t h slight subapical swelling; only 0.18-0.27
x BL. ANT PT/BASE 3.0-4.2 . . . . . . . . . . . . . . . . . Sitobion phyllanthi
SIPH not constricted at apex, and usually more than 0.27 x BL. ANT
PT/BASE 4.2-7.4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 R IV+V 0.65-0.80 X HT II. PT 1.4-2.0 X ANT III . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion ?mimosae
- R IV+V 0.85-1.05 x HT II. PT 1.0-1.4 x ANT I I I . . . . . . . . . . . . . . 8
8 Lateral abdominal tubercles absent. Dorsal abdomen of aptera unpig-
mented. CAUDA pale . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion takahashii
Lateral tubercles variably present on ABD TERG 3-5. Dorsal abdomen
of aptera often with dark transverse bands. CAUDA dusky . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion africanum group
9 CAUDA black with 8-16 hairs. Hind tibia with short, peg-like hairs in
a row on ventral side of basal half . . . . . . . . . . . . . . . . . Aphis eugeniae
CAUDA pale, dusky or black; if black then with 4-7 hairs, and hind tibia
without any peg-like hairs . . . . go to key to polyphagous aphids, p. 532
268 Phyllostachys

PHYLLOSTACHYS Gramineae

Phyllostachys seems to have a much smaller aphid fauna than other genera
of bamboos, but it may nevertheless be worthwhile to consult the keys to
Arundinaria and Bambusa for specimens that do not fit the species listed here.

Astegopteryx bambucifoliae
Ceratoglyphina bambusae
Glyphinaphis bambusae
Melanaphis bambusae
Paracolopha morrisoni
Phyllaphoides bambusicola
Takecallis arundicolens,
arundinariae, sasae, taiwanus

Key to aphids on Phyllostachys

1 Antenna of aptera much less than 0.5 x BL, and alata with very narrow
annular rhinaria extending more than half-way around segment . . . . . 2
Antenna of aptera more than 0.5 x BL or, if only alatae are present, these
have antennae bearing circular or oval secondary rhinaria . . . . . . . . . 5
2 Aptera w i t h o u t frontal horns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Aptera with frontal horns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Aptera (on roots) with antennae very short, hardly longer than the
elongate R IV+V. Tarsi 1-segmented. Body hairs fine . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracolopha morrisoni
Aptera (on leaves) with antennae short, but much longer than short
R IV+V. Tarsi 2-segmented. Body hairs thick and spine-like . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glyphinaphis bambusae
4 CAUDA and anal plate rounded. Frontal horns of aptera longer than
ANT I + II . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratoglyphina bambusae
CAUDA knobbed, anal plate bilobed. Frontal horns not longer than ANT
I + II . . . . . . . . . . . . . . . . . . Astegopteryx spp.; see Bambusa key, p. 83
5 CAUDA tongue-like, without a constriction, anal plate entire. ANT
PT/BASE more than 4. Antenna of alata with round secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis bambusae
- CAUDA knobbed, anal plate bilobed. ANT PT/BASE about 1. Antenna
of alata with transversely oval secondary rhinaria . . . . . . . . . . . . . . . . 6
6 Eyes without ocular tubercles. SIPH cones very flat, mammiform . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phyllaphoides bambusicola
 Picea 269

Eyes with ocular tubercles. SIPH cones truncate . . . . . Takecallis spp.

(for separation see Bambusa key)

PICEA Spruces Pinaceae

Many of the 38 species of spruce-feeding Cinara have only been found on one
or two hosts, but the more widely-distributed and/or more common species
show little evidence of specificity to particular species of Picea, so a single,
rather long, key is required, of which couplets 36 onwards deal with this genus;
only the apterous viviparae are keyed. Many of the North American Cinara
species are inadequately known, and their separation often relies heavily on
the original descriptions. Cinara nepticula Hottes and Cinara yukona Hottes
are only known from the alate morph so cannot be included in the key. Several
other separations, especially those involving species not seen by the authors
(marked *), are probably unreliable because they are based on too few
specimens.
Picea is the primary host for Adelgidae. The taxonomy of adelgids is
difficult because of their complex polymorphism (see Fig. 1). Identification of
adelgids on spruce requires examination of immatures of the fundatrix genera-
tion (usually found overwintering in or near the buds) and/or the alate migrant
(gallicola) that emerges from the gall the following summer. Even then, species
limits are often uncertain (most notably in the laricis group). However, there
is some degree of host specificity in Adelgidae, so it should be possible to
narrow down the possibilities considerably by using the key in conjunction
with the host plant list below and the descriptions of galls, etc. in the text.
The parthenogenetic generations of species in a number of genera of
Pemphiginae in the tribe Prociphilini, Pachypappa, Pachypappella, Pro-
ciphilus, live in white wax on the roots of Picea spp. as their secondary hosts.
The couplets (29-35) separating apterae of Pachypappa and Pachypappella on
spruce roots are based on the work of Danielsson (1990a). The root-feeding
morphs of Prociphilus (subgenus Stagona) cannot at present be satisfactorily
keyed to species.

Host Plant List

Picea abies (= excelsa) Adeiges abietis, lapponicus,
(Norway Spruce) lariciatus, laricis, pectinatae,
tardoides, viridis
Cinara braggii, costata, ezoana,
fornacula, piceae, piceicola,
pilicornis, pinicola, pruinosa
Elatobium abietinum
Pachypappa populi, tremulae,
vesicalis
270 Picea

Pachypappella lactea
Pineus cembrae, konowashiyai,
pineoides
Prociphilus (Stagona) xylostei
Tsugaphis sorini ssp. piceicola
P. asperata Cinara alba, piceae
(Chinese Spruce)
P. brachytyla Adelges glandulae
P. engelmanni Adelges cooleyi
Cinara braggii, coloradensis,
?curvipes, engelmanniensis,
fornacula, glehna, hottesi,
nimbata, obscura, pruinosa,
saskensis, vandykei
Elatobium abietinum
Pineus boycei, engelmanni,
pinifoliae
Prociphilus (Stagona) xylostei
P. glauca Adelges abietis, cooleyi, lariciatus,
(White Spruce) pectinatae ssp. ishiharai, tardus
Cinara acadiana, atripes, ?bonica,
?bonita, braggii, coloradensis,
costata, ?curvipes, fornacula,
hottesi, jucunda, nigripes,
obscura, pallidipes, piceae,
piceicola, pilicornis, pinicola,
pruinosa, saskensis, soplada,
thatcheri, vandykei, [yukona]
Essigella alyeska
Mindarus obliquus
Pachypappa rosettei, vesicalis
Pineus pinifoliae
Prociphilus (Stagona) xylostei
Tsugaphis sorini ssp. piceicola
P. glehnii Adelges pectinatae ssp. ishiharai
(Sakhalin Spruce) Cinara costata, ezoana, glehna,
nopporoensis, piceae, pinicola
Pineus cembrae
Prociphilus (Stagona) konoi
P. jezoensis (= ajanensis) Adelges abietis, japonicus,
(Yeddo Spruce) karafutonis, lapponicus,
pectinatae ssp. ishiharai
Cinara costata, ezoana, horii,
?piceae, pruinosa
 Picea 271

Elatobium momii/piceana
Pineus orientalis
Tsugaphis sorini ssp. piceicola
P. jezoensis var. hondoensis Adelges isedakii, torii
P. koraiensis Adelges laricis, pectinatae, viridis
Tsugaphis sorini ssp. piceae
P. koyamai Adelges abietis
P. likiangensis Adelges glandulae
Pineus sichunanus
P. mariana Adelges cooleyi, lapponicus,
(Black Spruce) lariciatus, lands
Cinara abietis, braggii, fornacula,
hottesi, mariana, nigripes,
nimbata, pruinosa, rara
Pachypappa rosettei, ?sacculi
Pineus floccus, pinifoliae, strobi
P. morrisonicola [Prociphilus formosanus]
P. obovata Adelges lapponicus, lands,
(Siberian Spruce) pectinatae, tardus
Cinara piceicola
Pineus cembrae, pineoides
[Prociphilus oriens]
P. omorika Adelges nordmannianae, prelli
Pineus pineoides
P. orientalis Adelges lands, merkeri,
nordmannianae, prelli, viridis
Cinara pilicornis, pruinosa
Pachypappa tremulae
Pineus orientalis
P. polita Adelges funitecta
(Tigertail Spruce) Pineus orientalis
P. pungens Adelges cooleyi, lapponicus,
(Colorado Spruce) lariciatus, lands, tardus
Cinara braggii, coloradensis,
costata, fornacula, glehna,
jucunda, piceae, pilicornis,
pruinosa, [wanepae]
Elatobium abietinum
Pachypappa ?sacculi, tremulae
Pineus pinifoliae, similis
Protohormaphis piceae
Tsugaphis sorini ssp. piceicola
272 Picea

P. purpurea Pineus sichunanus

P. retroflexa Adelges laricis ssp. pontaninilaricis
P. rubens Cinara braggii, fornacula,
(Red Spruce) nepticula, pinicola
Pineus floccus, pineoides,
pinifoliae
Prociphilus (Stagona)
piceaerubensis
P. schrenkiana Adelges lapponicus
Cinara piceae, pruinosa
Pineus pineoides
Tsugaphis sorini ssp. piceicola
P. sitchensis Adelges abietis, cooleyi, japonicus,
(Sitka Spruce) laricis, pedinatae ssp. ishiharai,
viridis
Aphis fabae
Aulacorthum (Neomyzus)
circumflexum
Cinara braggii, coloradensis,
costata, fornacula, gladalis,
nigripes pilicornis, pinicola,
sitchensis, vandykei
Elatobium abietinum
Mindarus obliquus
Pachypappa tremulae, vesicalis
Pachypappella lactea
Pineus pinifoliae, similis
Prociphilus (Stagona) xylostei
P. smithiana Adelges knucheli
(Himalayan Spruce) Cinara comata
Picea sp. Cinara alaskana, caudella

Key to spruce-feeding aphids

1 Abdomen of adults with a chitinous ovipositor (parthenogenetic females
oviparous). Apterae very small (less than 1.2mm) with greatly reduced
antennae of at most 3 segments. Alatae with short 5-segmented antennae
without secondary rhinaria but with a single large rhinarium on each of
ANT III, IV and V (Adelgidae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- No chitinous ovipositor (parthenogenetic females viviparous). Antennae
of apterae with at least 4 segments. Alatae (where known) with 5- or
6-segmented antennae with secondary rhinaria at least on ANT III . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
 Picea 273

2 Abdomen (all morphs) with only 4 evident pairs of spiracles, on ABD

TERG 2-5 (Pineus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Abdomen (all morphs) with 5 evident pairs of spiracles, on ABD TERG
2-6 (Adelges) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3 Adult aptera very small (BL about 0.8mm or less), dorsoventrally flat-
tened, free-living all year around on lignified tissue (in cracks of bark,
under scales, etc.). Head and prothorax fused into a well-sclerotized
shield, with conspicuous wax gland facets (Fig. 64A). (Alate morphs
unknown) . . . . . . . . . . . . . . . . . . . . . . . Pineus pineoides (/konowashiyai)
- Adult aptera with BL usually over 1 mm, found in or near galls on shoots
(usually fundatrices). Head and prothorax only lightly sclerotized, fused
or separate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Head of alata emerging from gall (gallicola) without wax pore plates
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus pinifoliae
- Head of alate gallicola with wax pore plates . . . . . . . . . . . . . . . . . . . . . 5
5 (Alate gallicola) Dorsal abdominal wax glands comprising separate small
groups, each with a few facets, distributed in irregular tranverse rows
between narrow crescent-shaped marginal wax pore plates (Fig. 64B)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus boycei*
- (Alate gallicola) Dorsal abdominal wax glands where present grouped in
spinal, or spinal and pleural, plates forming longitudinal (but often
incomplete) rows between the marginal plates, which are oval or broadly
crescent-shaped (e.g. Fig. 64C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 (Alate gallicola) Primary rhinarium on ANT V clearly extending more
than half-way around segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- (Alate gallicola) Primary rhinarium on ANT V not extending more than
half-way around segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 (Alate gallicola) Primary rhinarium on ANT V very large and elongate,
occupying more than 0.6 of total length of ANT V (Fig. 64D) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus floccus*
- (Alate gallicola) Primary rhinarium on ANT V not elongate, not usually
occupying more than 0.5 of total length of ANT V (Fig. 64E) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus orientalis
8 Abdomen of overwintering immature 'fundatrix' (pseudofundatrix) and of
adult aptera from gall (pseudofundatrix or apterous gallicola) almost
devoid of spinopleural wax glands posterior to ABD TERG 1, and with
only small marginal glands . . . . . . . . . . . . . . . . . . . . . . . . . Pineus similis
- Abdomen of overwintering immature fundatrix with spinal and pleural
wax glands on all tergites, and adult fundatrix with spinopleural glands
on ABD TERG 1-3, 1-4 or 1-5, and well-developed marginal glands on
ABD TERG 1-7 . . . . . . . . . . . . . . . . . . . Pineus cembrae (or sichunanus)
274 Picea
 Picea 215

9 Wax glands of overwintering immature fundatrix with simple, thin-walled

facets secreting solid wax filaments. Galls like miniature pineapples with-
out needles, hair or waxy bloom . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Wax glands of overwintering immature fundatrix (or pseudofundatrix)
with double-walled ring-like facets secreting hollow tubes of wax (e.g.
Figs 64L-N). Galls usually with groups of needles arising from them,
and/or with hairs and/or a waxy bloom . . . . . . . . . . . . . . . . . . . . . . . 14
10 Immature fundatrix with pleural as well as spinal and marginal sclerites
of thorax and abdomen bearing wax glands (i.e. 6 longitudinal rows of
wax pore plates). Alate gallicola with wax glands on head and prothorax
having many distinct facets with clear lumina . . . . . . . . . . . . . . . . . . . 11
- Immature fundatrix without wax glands on pleural sclerites of thorax and
abdomen; i.e. only spinal and marginal wax pore plates on each segment
(the spinal wax pore plates being on the inner margins of the sclerites).
Alate gallicola with wax glands on head and prothorax of granular appear-
ance, without facets or with a few indistinct facets . . . . . . . . . . . . . . 12
11 Immature fundatrix with apical hair on antenna much less than half length
of ANT III; wax gland facets situated around margins of spinal and
pleural sclerites of meso- and metathorax and ABD TERG 1-5 leaving
reticulate central areas. Alate gallicola with rhinaria on ANT I I I , IV and
V occupying more than half the length of their respective segments . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges pectinatae
- Immature fundatrix with apical hair on antenna about half of length of
ANT III; wax gland facets situated centrally in groups on spinal and
pleural sclerites of meso- and metathorax and ABD TERG 1-5. Alate
gallicola with rhinaria on ANT III, IV and V occupying less than half the
length of their respective segments . . . . . . . . . . . . . . . . Adelges knucheli
12 Immature fundatrix with wax pore plates on inner margins of meso- and
metathoracic spinal sclerites containing numerous small, mostly rounded,
pits, arranged in 2-6 areas of rather rounded shape (Fig. 64I). Total
number of pits in the most central areas only (Fig. 64H, labelled) of the
spinal wax pore plates of the meso- and metathorax plus ABD TERG 1-3
(i.e. a total of 10 central areas) is 64-94 . . . . . . Adelges nordmannianae

Fig. 64. A, adult aptera of Pineus pineoides, dorsal (right) and ventral views; dorsal abdominal wax glands
of alate gallicola of B, Pineus boycei (after Annand, 1928) and C, P. orientalis; antenna of alate gallicola of
D, Pineus floccus (after Annand, 1928), E, P. orientalis, F, Adelges laricis, G, A. cooleyi; H, overwintering
fundatrix larva of Adelges (subgenus Dreyfusia) showing positions of wax glands on spinal sclerites, with
the 'central areas' arrowed (total number of pits in these on both sides is counted); meso- and metathoracic
spinal sclerites (on left side only), showing shape and arrangements of wax glands in overwintering fundatrix
larva of I, Adelges nordmannianae, J, A. merkeri and K, A. prelli (all after Eichhorn, 1956); L, pattern of
wax glands on thoracic sclerites of overwintering fundatrix larva of Adelges japonicus (after Inouye, 1953);
wax glands on prothorax of overwintering fundatrix larva of M, A. abietis and N, A. viridis.
276 Picea

- Immature fundatrix with wax pore plates on inner margins of meso- and
metathoracic spinal sclerites divided into 3-4 often rather angular areas,
containing relatively large pits of irregular shape (Figs 64J, K). Total
number of pits in the most central areas only of the spinal wax pore plates
of meso- and metathorax plus ABD TERG 1-3 (10 central areas) is 15-56
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Immature fundatrix with total number of pits in 10 central areas (as
defined in couplet 12) usually more than 40 (range 34-56, mean about 45)
(Fig. 64J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges merkeri

- Immature fundatrix with total number of pits in 10 central areas usually

less than 40 (range 15-47, mean about 26) (Fig. 64K) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges prelli
14 Overwintering immature fundatrix or pseudofundatrix (where known)
with dorsal sclerites bearing single rather large ring-like wax pores, ringed
by several small less distinct glands of irregular shape (Fig. 64L). Alate
gallicola often but not always having ANT IV and V with broad, indistinct
basal articulation, 2 or more x wider than basal articular diameter of
ANT I I I (e.g. Fig. 64F) . . . . . . . . . . . . . . . . . . . . . . Adelges laricis group
(isedakii, japonicus, lapponicus, lariciatus, laricis, pontaninilaricis)
- Overwintering immature fundatrix (or pseudofundatrix) with dorsal
sclerites bearing groups of similar-sized ring-like wax pores. Alate
gallicola with basal articulation of ANT IV and V distinct and not much
broader than that of ANT III (Fig. 64G) . . . . . . . . . . . . . . . . . . . . . . . 15
15 Immature fundatrix (or pseudofundatrix) with dorsal sclerites bearing
groups of 8-30 ring-like wax pores. Wax glands on head and prothorax
of alate gallicola with large distinct facets . . . . . . . . . . . . . . . . . . . . . . 16
- Immature fundatrix with dorsal sclerites bearing groups of 2-7 ring-like
wax pores (e.g. Fig. 64M). Wax glands on head and prothorax of alate
gallicola absent or indistinct, without any clear facets . . . . . . . . . . . . 17
16 Rhinaria on ANT III, IV and V of alate gallicola transverse, slit-shaped,
that on V occupying less than 0.3 of length of segment (excluding short,
blunt PT) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges cooleyi
- Rhinaria on ANT III, IV and V of alate gallicola oval, that on V elongate
and occupying about 0.5 of length of segment (excluding PT, which is
pointed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges glandulae*
17 Alate gallicola with oblique vein of hindwing very short, almost straight,
at right angles to longitudinal vein, and weakly pigmented; rhinarium on
ANT V occupying less than 0.5 of total length of segment . . . . . . . 18
- Alate gallicola with oblique vein of hindwing either curved or bent
distally, with or without pigment; rhinarium on ANT V occupying about
0.5 of total length of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
 Picea 211

18 Overwintering 1st instar fundatrix (pseudofundatrix) elongate, BL about

2.5 x greatest body width, with pleurospinal sclerites of prothorax clearly
longer than broad (Fig. 64M). Nymph of alate gallicola (in gall) with 0-10
wax pores on each marginal sclerite of mesothorax . . . . . Adelges abietis
- Overwintering first instar fundatrix with BL about 1.7 x greatest body
width, and with pleurospinal sclerites of prothorax almost square
(Fig. 64N). Nymph of alate gallicola with 15-30 wax pores on each meso-
thoracic marginal sclerite . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges viridis
19 Alate gallicola with oblique vein of hindwing arising at right angles from
longitudinal vein and curved outward distally (not heavily pigmented?).
BL about 1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . Adelges karafutonis*
- Alate gallicola with oblique vein of hindwing heavily pigmented, arising
at an obtuse angle from longitudinal vein and mainly straight, with an
angular bend outward distally. BL 1.8-2.4 mm . . . . . . . . . Adelges torii
20 ANT PT/BASE more than 1. SIPH tubular, CAUDA tongue- or finger-
like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- ANT PT/BASE less than 1. SIPH on broad pigmented cones, or pore-
like, or absent. CAUDA short; rounded, triangular or knobbed . . . . 24
21 SIPH and CAUDA black. Hairs on legs long and fine, and those on
antennae more than 0.5 x basal diameter of ANT I I I . . . . . Aphis fabae
- SIPH and CAUDA pale or dusky. Hairs on legs and antennae short; those
on antennae much less than 0.5 x basal diameter of ANT I I I . . . . . . 22
22 Aptera with a dark horseshoe-shaped dorsal abdominal patch (Fig. 121I),
and alata with a solid dark dorsal abdominal patch . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum (Neomyzus) circumflexum
- Aptera and alata both without dark dorsal abdominal markings . . . . 23
23 ANT PT/BASE no more than 1.6 . . . . . . . . . . . . Elatobium abietinum
- ANT PT/BASE more than 2 . . . . . . . . . . . . . . . . . . . . Elatobium momii
24 SIPH either absent or present as pores, sometimes elevated, but never with
broad pigmented conical bases. R V not clearly distinct from R IV, or not
elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
- SIPH as large pores, usually placed on broad pigmented hairy cones
(Cinara spp.). R V distinct from R IV and elongate, 2 or more x its basal
width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
25 Dorsum entirely sclerotized, with head and prothorax fused. Antenna
4- or 5-segmented. CAUDA knobbed . . . . . . . . . . . . . . . . . . . . . . . . . . 26
- Dorsum mainly membranous, head and prothorax separate or, if fused,
then antennae are 6-segmented. CAUDA rounded or triangular . . . 27
278 Picea

Fig. 65. Dorsal (right) and ventral aspects of apterous vivipara of A, Protohormaphis piceae and
B, Tsugaphis sorini piceicola (both after Shaposhnikov and Gabrid, 1987).

26 Body of aptera broadly elliptical, greatly flattened and expanded laterally,

divided dorsally into cephalothorax and abdomen of approximately equal
size, no other segmentation being visible (Fig. 65B). Short legs and
4-segmented antennae hidden under body . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tsugaphis sorini piceicola
- Body of aptera of more normal aphid form (Fig. 65A). Antennae
5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protohormaphis piceae
27 Aptera with compound eyes and elongate, spindle-shaped body, BL more
than 2 x maximum body width. Wax glands absent . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Essigella alyeska
- Aptera with reduced eyes (3-7 facets), BL less than 2 x maximum body
width. Wax glands well developed, at least on abdomen of aptera . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 Aptera with R IV+V about 0.5 x ANT BASE VI, and with marginal wax
pore plates present on ABD TERG 1-8 or 2-8; spinal and pleural wax pore
plates reduced or absent except on most posterior tergites. Alata with
media of forewing usually once-branched (on needles) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mindarus obliquus
 Picea 279

Fig. 66. Fore tarsus of apterous exule of A, Pachypappa tremulae, B, Pachypappella lactea and
C, Pachypappa vesicalis.

- Aptera with R IV+V about 1.0 x ANT BASE VI or longer, and with
marginal wax pore plates reduced or absent, at least on ABD TERG 3-8;
large spinal and pleural wax pore plates on ABD TERG 4-7. Alata
(sexupara) with media of forewing usually unbranched (on roots) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 R IV+V without a pale subapical zone. Aptera with thick, spine-like hairs
on HT I, similar in size to those at apex of tibia. Alate sexupara with
secondary rhinaria on ANT V . . . . . . . . . . . Prociphilus (Stagona) spp.
- R IV+V with a distinct pale subapical zone. Aptera with hairs on HT I
fine or spine-like, if spine-like then often smaller than those at apex of
tibia. Alate sexupara without secondary rhinaria on ANT V . . . . . . 30
30 (Remaining couplets in this part of key refer only to aptera.) Hairs on HT
I very small and thin, or if thick basally then with finely-pointed apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
- Hairs on HT I thick and sometimes spine-like, with short or blunt apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
31 Segments I and II of all tarsi distinctly separated from each other. Seg-
ment I of fore tarsus with 2-3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
- Segments I and II of tarsi not distinctly separated. Segment I of fore tarsus
with 2 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
32 Hairs on HT I and apical dorsal hairs on HT II very small, 4-8 µm long,
with abrupt apices. Segment I of fore tarsus usually with 2 hairs; if a third
middle hair is present it is longer than the lateral hairs . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa rosettei*
- Hairs on HT I and apical dorsal hairs on HT II longer, 10-15 µm, with
finely-pointed apices. Segment I of fore tarsus almost always with 3 hairs,
the middle hair shorter and blunter than the lateral hairs (Fig. 66A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa tremulae
280 Picea

Fig. 67. Hairs on outer side of hind tibia of aptera of A, Cinara piceae, B, C. piceicola and C, C. pilicornis.

33 ANT PT only 0.015-0.02 mm long. Legs short, with length of hind femur
less than 4 x its maximum width. All tarsi reduced, with segments I and
II not distinctly separated (HT II less than 0.1 mm long). Second tarsal
segments with dorsal apical hairs short and spine-like (Fig. 66B) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappella lactea
- ANT PT more than 0.02 mm long. Legs longer, with hind femur more
than 4 x its maximum width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34 Antenna usually 6-segmented, with PT finger-like, 0.034-0.05 mm long.
Hairs on first tarsal segments spine-like and similar in size to those at
apices of tibiae. Hind tibia on dorsal side bearing 2-5 spine-like hairs with
short, blunt apices . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa populi*
- Antenna usually 5-segmented, with PT less than 0.035 mm long. Hairs on
first tarsal segments distinctly shorter than those at apices of tibiae (e.g.
Fig. 66C). Hind tibia with some hairs on dorsal side with long, pointed
apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35 Hairs arising from dorsal abdominal wax pore plates are more than 25 µm
long. Each wax gland facet with a dark circle inside a pale outer ring
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa vesicalis
- Hairs arising from abdominal wax pore plates are less than 25 µm long.
Wax gland facets without dark centres . . . . . . . . ?Pachypappa sacculi*
36 (Remaining couplets all refer only to apterous viviparae.) Hairs on outer
side of hind tibia short and rather thick (e.g. Fig. 67A), only more than
0.06mm long in very large aphids, mostly less than or not exceeding
0.5 x width of hind tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . 37
- Hairs on outer side of hind tibia more than 0.06 mm long and much more
than 0.5 x width of hind tibia at midpoint . . . . . . . . . . . . . . . . . . . . . 41
37 Large rotund aphid, BL 3.2-6.7 mm long, but usually more than 4.5 mm.
Coxae black. R IV more than 0.30mm . . . . . . . . . . . . . . Cinara piceae
 Picea 281

- BL 2.4-4.2 mm. Coxae (where colour known) light brown. R IV less than
0.25 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 PT less than 0.03 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara jucunda*
- PT more than 0.04 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
39 R IV only about 0.17 mm (in specimen of BL about 2.5 mm) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara caudelli*
- R IV more than 0.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
40 HT II about 0.29mm (in specimen of BL 3.1 mm). Hairs on SIPH cones
rather evenly distributed over entire pigmented area . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara soplada*
- HT II 0.33 mm or more. Hairs on SIPH cones concentrated on apical half
around pore, with basal part of pigmented area having only sparse hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara coloradensis
41 Hairs on outer side of middle section of hind tibia all less than 0.12mm
long (e.g. Fig.67B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
- All or many of hairs on outer side of middle section of hind tibia of length
exceeding 0.12 mm (e.g. Fig. 67C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
42 Hind tibiae uniformly dark brown to black . . . . . . . . . Cinara atripes*
- Hind tibiae with at least the basal or middle section paler . . . . . . . . 43
43 SIPH cones pale, almost concolorous with dorsal cuticle . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara fornacula
- SIPH cones pigmented, much darker than rest of dorsal cuticle . . . 44
44 HT II more than 0.32 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
- HT II less than 0.32 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
45 BL less than 4.6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara piceicola
- BL more than 5.0mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
46 R IV about 0.22mm. Antennae and legs short; ANT III less than
0.1 x BL, hind tibia less than 0.4 x BL . . . . . . . . . . . . . Cinara bonita*
- R IV 0.40-0.44 mm. Antennae and legs much longer; ANT III more than
0.15 x BL, hind tibia more than 0.6 x BL . . . . . . . . Cinara saskensis*
47 Hind tibia more than 1.65 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
- Hind tibia less than 1.65 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
48 R IV+V about 0.42 mm . . . . . . . . . . . . . . . . . . . . . . . . . Cinara acadiana*
- R IV+V 0.36 mm or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
49 R IV+V 0.24mm or less. ANT III with 1-2 secondary rhinaria distally
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara vandykei
282 Picea

- R IV+V 0.24-0.36 mm. ANT III without secondary rhinaria . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara hottesi
50 ANT VI (including PT) 0.15-0.18 mm, as long as or a little longer than
ANT V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara sitchensis
- ANT VI (including PT) 0.12-0.14 mm, a little shorter than ANT V . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara obscura/pallidipes
51 ABD TERG II-VI each with a pair of large dark patches, forming two
longitudinal rows on dorsum . . . . . . . . . . . . . . . Cinara engelmanniensis
- ABD TERG II-VI without paired dark patches (except around siphunculi)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
52 Hind tibiae uniformly dark brown or black . . . . . . . . . . . . . . . . . . . . . 53
- Hind tibiae pale, or at least paler basally or medially than at apex . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
53 SIPH cones large and dark . . . . . . . . . . . . . . . . . . . . . . . . Cinara nigripes
- SIPH cones very small and pale . . . . . . . . . . . . . . . . . . . . . . Cinara horii
54 ANT IV without any rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
- ANT IV with at least one rhinarium . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
55 Fore tibiae black, contrasting with pale mid- and hind tibiae. SIPH cones
pale. Hairs on antennae long (up to 160 µm), and sparse . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara atroalbipes
- Fore tibiae similarly pigmented to mid- and hind tibiae. SIPH cones dark.
Hairs on antennae about 120µm long, numerous . . . . Cinara bonica*
56 HT II longer than maximum diameter of SIPH cones, which are small and
often rather pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pilicornis
- HT II shorter than maximum diameter of SIPH cones . . . . . . . . . . . 57
57 Hind femora either wholly pale or with patchy or banded pigmentation.
Tibial hairs dark, often with pigmented bases. Dorsal abdominal hairs
often arising from small scleroites. (Alata with media of forewing once-
branched) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
- Hind femora sometimes wholly pale but usually pale basally becoming
dark distally. Tibial hairs pale or dusky, usually with unpigmented bases,
and dorsal abdominal hairs rarely arising from small scleroites. (Alata
with media of forewing twice-branched) . . . . . . . . . . . . . . . . . . . . . . . . 60
58 R IV+V as long as or longer than HT II. (Alata with forewings marked
with fuscous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara costata
- R IV+V shorter than HT II. (Forewings of alata with or without fuscous)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
 Pinus 283

59 Longest hairs on ANT III about one quarter of length of ANT III. (Alata
with forewings marked with fuscous) . . . . . . . . . . . . . . . Cinara nimbata
- Longest hairs on ANT III about one half of length of ANT III. (Alata
with hyaline forewings) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara comata
60 R IV+V 1.1-1.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pruinosa
- R IV+V less than 1.1 x HT II ................................ 61
61 ANT III 3.4-4.1 x longer than the longest hair borne upon it. HT II
0.30-0.48 mm long, 1.1-1.4 x R IV+V. Coxae, femora and tibiae all
rather pale; tibiae only dark at apices . . . . . . . . . . . . . . . Cinara braggii
- ANT III 2.4-3.1 x the longest hair borne upon it. HT II 0.28-0.32 mm
long, 1.0-1.2 x R IV+V. Legs darker, with tibiae dark on about distal
third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
62 ANT III more than 0.15 x BL, bearing hairs up to 160µm long . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara glacialis*
- ANT III 0.08-0.13 x BL, with longest hairs 90-130 µm long . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara glehna/mariana*

PILGERODENDRON Cupressaceae

Pilgerodendron uviferum Neophyllaphis michelbacheri

PINUS Pines Pinaceae

There are about 170 pine-feeding aphids, including more than 100 species of
Cinara. A single key including all the Cinara species would be unwieldy and
difficult to use. Fortunately there is a fairly well-established subgeneric classi-
fication of Pinus to which many (but by no means all) pine-feeding Cinara
species conform. We have used this subgeneric structure wherever possible to
key together Cinara species that feed on related pines. We have also made use
of it for separation of some closely related and morphologically similar species
of Essigella, a wholly North American genus of Lachnidae recently revised
using multivariate techniques by Sorensen (1992).
The main key that follows goes as far as possible with the identification
of all pine-feeding aphids to species level, except the genera Cinara and
Essigella. In the case of the adelgid genus Pineus, the secondary host morphs
(sistens, progrediens) on Pinus are very difficult to identify because of their
284 Pinus

lack of species-constant characters, and even the partial key provided should
be treated very warily.
The separate species of Pinus are then listed with the aphids recorded from
them, each followed where necessary by keys to Cinara and/or Essigella.
These keys can only be applied to apterous viviparae.

Key to pine-feeding aphids (apterous viviparae only, except where stated

otherwise)
1 Adults without chitinous ovipositor. BL usually more than 1.3 mm. Head
and prothorax separate, and antenna of at least 4 segments. (Parth-
enogenetic adults viviparous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Adults with chitinous ovipositor. BL less than 1.2mm. Head and pro-
thorax covered by a cephaloprothoracic shield. Antennae greatly reduced,
stump-like. (Parthenogenetic forms oviparous) . . . . . . . . . . . . . . . . . . 25
2 ANT PT/BASE more than 1. SIPH tubular. (Dorsal abdomen with a
dark, more-or-less horseshoe-shaped patch) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum (Neomyzus) circumflexum
- ANT PT/BASE less than 1. SIPH on broad pigmented cones, or pore-
like, or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Rostrum more than 1.5 x BL . . . . . . . . . . . . . . . . . . . . . . Stomaphis pini
- Rostrum less than 1.5 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Eyes reduced to triommatidia. (On roots) . . . . . . . . . . . . . . . . . . . . . . . 5
- Eyes multifaceted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5 Anal plate extended posteriorly as a pigmented tongue-like process, bear-
ing the bluntly conical CAUDA, which is longer than its basal width and
bears more than 15 hairs (Fig. 68A). R IV+V with 6-16 accessory hairs
............................................................. 6
- CAUDA rounded, broader than long, with 2-8 hairs (e.g. Fig. 68B).
R IV+V with 2-8 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 CAUDA with 40-60 hairs, R IV+V with 10-16 accessory hairs. (Alate sex-
upara with 20-30 secondary rhinaria on ANT III, 5-10 on IV and 9-15
on V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus carolinensis
- CAUDA with 20-35 hairs, R IV+V with 6-10 accessory hairs. (Alate
sexupara with 5-8 secondary rhinaria on ANT III, 1-4 on IV and 0 on
V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus piniradicivorus
7 R IV+V elongate, about 0.2 mm long, with 4-8 accessory hairs. (Alate sex-
upara with 3-9 secondary rhinaria on ANT VI BASE, and with
pterostigma ending bluntly well before tip of wing) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus caryae fitchii
- R IV+V less than 0.15 mm long, with 2-4 accessory hairs. (Alate sexupara
 Pinus 285

Fig. 68. A, CAUDA and anal plate of aptera of Prociphilus piniradicivorus; B, CAUDA of aptera of
P. pini; hind tibia of aptera of C, P. crataegistrobi and D, P. pini.

with 0-4 secondary rhinaria on ANT VI BASE, and with distal end of
pterostigma tapering, curved around tip of wing) . . . . . . . . . . . . . . . . . 8
8 R IV+V with 2 accessory hairs. Hind tibia thick, with 1-3 short, spine-like
hairs near middle on outer side (Fig. 68C). (Alate sexupara with 1-4 secon-
dary rhinaria on ANT VI BASE) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus (Stagona) crataegistrobi
- R IV+V with 3-4 accessory hairs. Hind tibia narrower, with several hairs
distributed along outer side which are fine-pointed, not distinctly spine-
like (Fig. 68D). (Alate sexupara without secondary rhinaria on ANT VI
BASE) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus (Stagona) pini
9 SIPH as large pores on usually pigmented hair-bearing cones . . . . . 10
- SIPH as small pores without associated hairs . . . . . . . . . . . . . . . . . . . 17
10 R V acutely pointed, dagger-shaped, usually twice or more as long as its
basal width (e.g. Fig. 69A). SIPH cones of aptera often large and with
numerous hairs . . . . . Cinara spp. (see separate keys in Host Plant List)
- R V short, flask-shaped, pointed only at tip, similar in length to or not
much longer than its basal width (e.g. Figs69B, C). SIPH cones always
with few hairs, in 1-3 rings around pore . . . . . . . . . . . . . . . . . . . . . . . 11
11 Hairs on ventral side of hind femur very short, thick and peg-like
(Fig. 70A). Hind legs wholly dark except for extreme base of femur, and
very elongate; total length about 2 x BL . . . . Schizolachnus flocculosus
- Hairs on ventral side of hind femur long, not peg-like. Hind legs long but
less than 2 x BL, with at least basal fifth of femur pale . . . . . . . . . 12
 286 Pinus

Fig. 69. R V of A, Cinara pini, B, Schizolachnus pineti, and C, S. obscurus.

Fig. 70. A, ventral side of hind femur of Schizolachnus flocculosus (aptera); B, middle section of hind
tibia of Schizolachnus curvispinosus (aptera); C, hair-bearing scleroites on ABD TERG 3 of
S. curvispinosus (aptera); ANT VI of D, S. pineti and E, S. parvus.

12 Hind tibia wholly dark bearing long, thick, curved hairs with blunt apices
(Fig. 70B). Dorsal abdominal hairs thick and spine-like, arising from
transverse rows of small sclerites (Fig. 70C) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus curvispinosus
- Hind tibia pale or dark, bearing mainly long, finely-pointed hairs. Dorsal
abdominal hairs also long and fine, not arising from sclerites ..... 13
13 Hind tibia more than 20 x longer than its width at midpoint ........
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus piniradiatae
 Pinus 287

- Hind tibia less than 20 x longer than its width at midpoint . . . . . . 14

14 R V very short and stumpy with short tip (Fig. 69B); less than 46 µm long
from base to tip, and less than 0.45 x R IV . . . . . . . . . . . . . . . . . . . . 15
- R V with long tip (e.g. Fig. 69C); more than 46 µm long and more than
0.45 x R IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15 Hind tibia pale or dark, very densely hairy (about 250 hairs per mm). PT
very short; ANT PT/BASE less than 0.25 (Fig. 70D). (Alata with once-
branched media and without secondary rhinaria on ANT IV) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus pineti
- Hind tibia dark and less densely hairy (about 150 hairs per mm). ANT
PT/BASE more than 0.25 (Fig. 70E). (Alata with unbranched media and
1 or more secondary rhinaria on ANT IV) . . . . . Schizolachnus parvus
16 Hind tibia pale. (Alata with unbranched media) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus orientalis
- Hind tibia dark. (Alata with usually once-branched media) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizolachnus obscurus
17 Antenna 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
- Antenna 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
18 Hairs on ABD TERG 1-6 not arising from sclerites . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus pumilae
- Hairs on ABD TERG 1-6 arising from small dark rounded sclerites
(scleroites) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 ANT III less than 0.24mm long and bearing hairs less than 20 µm long.
Hairs on ABD TERG 1-7 short and mostly pointed, 4-25 µm long, usually
shorter than the diameter of their basal scleroites . . . . . . . . . . . . . . . 20
- ANT III more than 0.25 mm long and bearing hairs up to 40-130 µm long.
Hairs on ABD TERG 1-7 usually longer than or as long as the minimal
width of their basal scleroites; if short and pointed on more anterior
tergites, then at least those on ABD TERG 7 are more than 25 µm long,
dark and with blunt or furcate apices . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20 Capitate hairs on dorsal side of hind tibia often longer than width of tibia
at midpoint. Hair-bearing scleroites on ABD TERG 1-5 in a single
transverse row on each tergite (Fig. 71 A) . . . . . . . . Eulachnus nigricola
- Capitate hairs on dorsal side of hind tibia not exceeding width of tibia
at midpoint. Hair-bearing scleroites on ABD TERG 1-5 placed irregu-
larly, not in a single transverse row (Fig. 7IB) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus brevipilosa
21 Longest hairs on ANT III 20-50 µm long, rarely more than one hair over
40µm. ABD TERG 1-3 with longest hairs 15-40µm long, pointed, and
hardly pigmented (Fig. 71C) . . . . . . . . . . Eulachnus tuberculostemmata
288 Pinus

Fig. 71. Arrangement of spinopleural hair-bearing scleroites between muscle plates on ABD TERG 3:
A, Eulachnus nigricola; B, Eu. brevipilosa; C, Eu. tuberculostemmata; D, Eu. thunbergii; E, Eu. rileyi; f, Eu. agilis.
 Pinus 289

- Several of longest hairs on ANT I I I more than 40µm long. ABD TERG
1-3 with longest hairs 25-145 µm long, but if under 40 µm then they are
often blunt or slightly capitate and dark like those on ABD TERG 7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 Hair-bearing scleroites on ABD TERG 1-5 numerous, in two irregular
transverse rows on each tergite, the anterior row having more scleroites
than the posterior row (Fig. 71D). Fore femur dusky or dark and very
thick; its length less than 3.5 x its maximum width . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus thunbergii
- Hair-bearing scleroites on ABD TERG 1-5 often in two rows, but then
the anterior 'row' often has very few scleroites (often only 2), and never
has more than the posterior row. Fore femur pale or dark, more than
3.5 x longer than its maximum width . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 ABD TERG 3 with longest hairs 100-155 µm, usually with finely pointed
apices (Fig. 71E). ANT V usually clearly longer than (1.1-1.5 x) HT II.
Hind femur uniformly dark except at base; much darker than basal half
of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus rileyi
- ABD TERG 3 with longest hairs 25-110 µm, and usually with blunt apices
(Fig.71F). ANT V usually shorter than (0.6-1.2 x ) HT I I . Hind femur
pale or mottled, mainly about as dark as basal half of ANT I I I . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eulachnus agilis
24 Claws normal, with simple curved acute apices. ABD TERG 2-7 mem-
branous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudessigella brachychaeta
- Claws with modified apices; bifurcate, or double (Fig. 73). ABD TERG
2-7 lightly to heavily sclerotized (but not necessarily pigmented) . . . . .
. . . . . . . . . . . . . . . . Essigella spp. (see separate keys in Host Plant List)
25 Cephaloprothoracic shield (henceforth CPS) with sutures between head
and thorax and along midline. Wax glands on CPS comprising mainly
separate, large facets that are distinctly subdivided (Fig. 72A) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus pinifoliae
- CPS entire. Wax glands on CPS consisting mainly of groups of facets that
are not individually subdivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Wax glands on anteriodorsal part of CPS (i.e. between eyes) mainly com-
prising groups of 2-10 facets that are often loosely grouped or at least
not closely pressed together, so that all except the largest retain a more-or-
less circular shape (e.g. Figs 72B, C) . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
- Wax glands between eyes mainly comprising more extensive groups or net-
works of 10 to more than 100 closely packed facets, mostly pressed
together so that they tend to be polygonal in shape (e.g. Figs. 72D, E) or
look like overlapping tiles when viewed obliquely . . . . . . . . . . . . . . . . 30
27 Wax glands between eyes comprising groups of 2-6 large facets that are
mostly much larger than the ocelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
290 Pinus

Fig. 72. Dorsal (right) and ventral aspects of A, Pineus pinifoliae (after Annand, 1928) and B, P. pini,
showing details of wax gland structure and (for pini) antenna; wax gland facets mesiad to ocelli of
C, P. boerneri, D, P. cembrae and E, P. strobi.

Fig. 73. Tarsal claws of Essigella californica.

 Pinus 291

- Wax glands between eyes mainly comprising groups of 2-10 smaller

facets, the largest of which are not larger than or only slightly larger than
the ocelli (maximum internal diameter about 10µm) . . . . . . . . . . . . . 29
28 Wax gland facets in groups on CPS contiguous . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus ghanii/wallichianae
- Wax gland facets on CPS in loose groups, not contiguous . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus harukawai* / cladogenous*
29 Antennae of summer progredientes (i.e. adult apterae collected at base of
1-year-old needles) usually reduced to a single segment and fused to CPS,
rarely articulated; total length usually less than 58 µm. Median wax pore
groups often present on two or more abdominal tergites. Minimum
diameter of smallest facet in wax plates between eyes usually more than
8 µm (Fig. 72C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus boerneri
- Antennae of summer progredientes (Fig. 72B) usually with 2-3 segments,
not fused to CPS, usually more than 58 µm long. Median wax pore groups
usually only on at most one abdominal tergite. Minimum diameter of
smallest facet in wax plates between eyes often less than 8 µm (Fig. 72B)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus pini/orientalis
30 Wax glands on CPS between eyes very extensive; groups closest to the
ocelli usually have more than 40 facets (Fig. 72D) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus cembrae/coloradensis
- Wax glands on CPS between eyes usually with less than 30 facets per
group (e.g. Fig. 72E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31 Wax gland facets between eyes about 2 x diameter of the ocelli and up
to 2-3 x the diameter of facets in the very small groups on the posterior
margin of the CPS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pineus sylvestris*
- Wax gland facets between eyes mostly similar in size to the ocelli and to
the largest facets in the well-developed groups on the posterior margin of
the CPS . . . . . . . . . . Pineus strobi/matsumurai/hosayai/?cortecicolus*

Host Plant List (with keys to apterae of Cinara species)

Pinus albicaulis Cinara anzai, apini, ferrisi,
(Whitebark Pine) inscripta, medispinosa,
murrayanae, oregoni
Essigella californica
Prociphilus caryae ?ssp.
arbutifoliae

Key to Cinara spp. on Pinus albicaulis

1 BL more than 4.5 mm. Tibiae wholly black. Length of sclerotized part of
stylet groove (Fig. 74A) more than 2.4 mm . . . . . . . . . . . . . . . . . . ferrisi
292 Pinus

Fig. 74. A, rostrum of Cinara showing measurement of sclerotized part of stylet groove; B, SIPH of
Cinara anzai (aptera); C, HT I of C. atra; 0, SIPH of C. pinona (aptera); hairs on dorsal side of hind tibia
of aptera of E, C. apacheca and F, C. edulis.

BL less than 4.5 mm. Tibiae with paler section on basal half. Length of
sclerotized part of stylet groove less than 2.4 mm . . . . . . . . . . . . . . . . . 2
2 Hairs on dorsal side of hind tibia only about 0.5 or less of width of tibia
at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Hairs on dorsal side of hind tibia of length similar to width of tibia at
midpoint, or longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 ABD TERG 1-7 each with a pair of irregularly-shaped or broken pig-
mented areas, of similar extent on all tergites . . . . . . . . . . . . inscripta*
At least ABD TERG 3-6 without pigmented sclerites . . . . . . . . . . apini
4 SIPH bases rather small (maximum diameter less than 0.35 mm) and very
asymmetrical, extending about twice as far anteriorly as posteriorly
(Fig. 74B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . anzai
SIPH bases large and fairly symmetrical; maximum diameter more than
0.4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 BL 4.0-4.4 mm. R V very short, 0.06 mm or less, only about one third
of length of R IV. HT II about 0.4mm . . . . . . . . . . . . . . . . . . oregoni*
- BL 2.6-3.8 mm. R V 0.08-0.10 mm, more than one third of length of
 Pinus 293

R IV. HT II less than 0.3 mm . . . . . . . . . . . . medispinosa/murrayanae

(for separation of these species see key under P. contorta)

P. armandii Cinara orientalis

P. attenuata Cinara essigi
(Knobcone Pine) Essigella californica, essigi
(Use key to Cinara spp. on pines of subsect. Oocarpae under P. oocarpa)
(The two Essigella species are separated in the key under P. ponderosa)
P. australis = palustris
P. austriaca = nigra
P. banksiana Cinara banksiana, brevispinosa,
(Jack Pine) canatra, cronartii, nigra,
ontarioensis, pergandei, pinea,
pini, piniradicis, pinivora,
russellae, watsoni
Essigella alyeska
Pineus boerneri, coloradensis,
havrylenkoi, pini
Schizolachnus piniradiatae
(For Cinara spp., use key to Pinus subsect. Contortae, under P. contorta)
P. brutia = halepensis var. brutia
P. canariensis Cinara atlantica, maghrebica,
(Canary Islands Pine) maritimae, ponderosae
Essigella californica
Eulachnus rileyi,
tuberculostemmata
Pineus boerneri

Key to Cinara spp. on Pinus canariensis

1 HT I elongate, 0.16-0.22 mm long, at least 0.5 X HT II (Fig. 76A). R IV
0.24-0.30 mm long, more than 0.5 x ANT III . . . . . . . . . . . . maritimae
- HT I less than 0.14mm long, less than 0.5 x HT II. R IV 0.12-0.22 mm
long, less than 0.5 x ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Maximum diameter of SIPH cone 0.21-0.37 mm, less than the length of
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maghrebica
Maximum diameter of SIPH cone 0.40-0.83 mm, about equal to or
greater than length of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ABD TERG 5 with less than 20 hairs between SIPH bases, the longest
10-70 µm long (Fig. 80F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae
- ABD TERG 5 with more than 20 hairs, 60-180 µm long, between SIPH
bases (Fig. 80E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
294 Pinus

P. caribaea Cinara atlantica, maritimae,

(Caribbean Pitch Pine) melaina, ponderosae
Essigella californica
Pineus boerneri
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
P. cembra Cinara cembrae, mongolica, pini
(Swiss Stone Pine) Eulachnus pumilae
Pineus cembrae

Key to Cinara spp. on Pinus cembra and other pines of subsect. Cembrae
(koraiensis, pumila, sibirica)
1 Hairs on ABD TERG 5 long (80-180µm). ANT III shorter than IV + V
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Hairs on ABD TERG 5 short (4-40 µm). ANT III as long as or longer
than IV + V together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Head, pronotum and mesonotum dark. Total antennal length about
0.45 x BL. ABD TERG 8 with about 30 hairs . . . . . . . . . . . mongolica
Head and thorax pale dorsally. Total antennal length about 0.35 x BL.
ABD TERG 8 with about 20 hairs . . . . . . . . . . . . . minoripinihabitans*
3 R IV 0.14-0.22 mm long, about 2 x R V. Hairs on ANT VI BASE max-
imally about 1.5 x basal diameter of segment . . . . . . . . . . . . . . . . . pini
- R IV 0.21-0.29 mm, more than 2 x R V. Hairs on ANT VI BASE
maximally about 2 or more x basal diameter of segment . . . . . . . . . 4
4 R IV less than 0.25 mm long, bearing 8-10 accessory hairs . . . . cembrae
R IV more than 0.25 mm long, bearing 14-22 accessory hairs . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . watanabei

P. cembroides Cinara tanneri, terminalis,

(Mexican Pinyon Pine) wahtolca
Essigella hoerneri
Eulachnus rileyi

Key to Cinara spp. on Pinus cembroides and other pines of subsect. Cem-
broides (edulis, monophylla, quadrifolia)
Parts of this key are based on Hottes' (1960b) key to Cinara species on
P. edulis. Most of the Cinara species described from pinyon pines are based
on inadequate material, and further study is likely to reveal synonymies.
l R IV more than 0.28mm long, with more than 25 accessory hairs . . . 2
- R IV less than 0.27 mm long, with less than 25 accessory hairs . . . . 3
 Pinus 295

2 R IV 0.5 mm or more long, much longer than ANT III, with more than
70 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . puerca
- R IV less than 0.4 mm long, shorter than ANT III, with less than 40 acces-
sory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . tanneri
3 Diameter of SIPH base 0.25 mm or less . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Diameter of SIPH base 0.27 mm or more . . . . . . . . . . . . . . . . . . . . . . . 5
4 BL 1.6-2.2 mm. Legs with contrasting pale and dark sections . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . caliente
- BL about 2.6mm. Legs dark, without pale sections . . . . . . . . . poketa*
5 Ventral hairs on HT I longer than maximum width of segment, with apices
attenuated, the more distal hairs often having their apices sharply crooked
(Fig.74C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atra
- Ventral hairs on HT I mostly not longer than maximum width of HT I,
or if slightly longer then without attenuated and crooked apices . . . . . . 6
6 Hind tibia densely clothed in hairs (more than c. 300 per mm), with those
on dorsal side mostly arising at an angle of less than 45° and curved so
that distally they are almost parallel to the tibia (Figs 74E, F) . . . . . 7
- Hind tibial hairs usually less dense, but if at a density of more than
c. 300 per mm then those on dorsal side arise at an angle of 45° or more
and with apices pointing away from tibia . . . . . . . . . . . . . . . . . . . . . . . 10
7 Hairs on hind tibia short and spine-like (Fig. 74E), maximally about 50 µm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . apacheca
- Hairs on hind tibia fine (e.g. Fig. 74F), maximally more than 50 µm long
............................................................. 8
8 SIPH cone uniformly pigmented, with maximum diameter 0.3-0.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . edulis
- SIPH cone two-toned, with only the upper, constricted, part deeply
pigmented (e.g. Fig. 74D); maximum diameter about 0.35mm . . . . . 9
9 Length of pale section on basal half of hind tibia less than one quarter
of total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinona
- Length of pale section on basal half of hind tibia more than one quarter
of total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . metalica*
10 Hairs on hind tibia small and fine, maximally about 85 µm long, with
finely pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Hairs on hind tibia longer or, if less than 85 µm long, then coarse with
blunt or abruptly acute apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Hairs on dorsal side of hind tibia mainly standing almost at right angles
to tibia. ANT VI BASE with less than 12 hairs . . . . . . . . . . . . rustica*
296 Pinus

- Hairs on dorsal side of hind tibia set at a more acute angle to tibia. ANT
VI BASE with more than 12 hairs . . . . . . . . . . . . . . . . . . . . . . . . . pinata
12 ANT III less than 0.33 mm long, much shorter than maximum diameter
of SIPH cone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nitidula
- ANT III more than 0.35 mm long, usually about as long as or slightly
longer than maximum diameter of SIPH cone . . . . . . . . . . . . . . . . . . 13
13 ABD TERG 5-7 (or 6-7) each with a single pair of small dark sclerites.
Hairs on hind tibia maximally 70-150 µm . . . . . . . . . . . . . . . . . wahtolca
- ABD TERG 5-7 with or without sclerites, but not each with a single pair.
Hairs on hind tibia maximally 50-80 µm . . . . . . ponderosae/terminalis

P. clausa Cinara atlantica, cronartii,

(Alabama Pine, Sand Pine) pergandei, pinivora, taedae,
walsoni
Pineus pini
(Use key to Cinara spp. on Pinus subsect. Contortae under P. contorta)
P. contorta (including var. latifolia Cinara, brevispinosa, canatra,
and var. murrayana) contortae, medispinosa,
(Lodgepole Pine, Shore Pine) murrayanae, nigra, oregonensis,
parvicornis, pergandei, pinea
Essigella braggii, californica,
critchfieldi, knowltoni
Pineus coloradensis, havrylenkoi,
pini
Schizolachnus parva, pineti,
piniradiatae

Key to Cinara spp. on Pinus contorta and other pines of subsect. Contortae
(banksiana, clausa, virginiana)
1 Dorsal abdomen almost completely covered by an extensive pigmented
area that is fused at least between tergites 4 and 6 and usually merges with
SIPH bases (Figs 75A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Dorsal abdomen with any dark markings divided segmentally and not
usually merged with SIPH bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Subgenital plate (of vivipara) with only 6-14 hairs. R IV with only 4-7
accessory hairs. SIPH with only 10-12 short hairs (maximally 40 µm)
....................................................... russellae
- Subgenital plate with 30-40 hairs. R IV with 8-16 accessory hairs. SIPH
with 20-35 longer hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ANT VI BASE with 10-15 hairs. R IV with 10-16 accessory hairs,
including several on side away from stylet groove . . . . . . . . . . . gracilis
 Pinus 297

- ANT VI BASE with 8-11 hairs. R IV with 7-11 accessory hairs in two
rows alongside stylet groove, rarely with 1-2 on opposite side . . . . . 4
4 Dorsal cephalic hairs rather sparse, situated mainly alongside dorsal
suture and near antennal bases, maximally 60 µm long, often shorter than
the distances between their bases. SIPH cones shallow with their bases
very extensive, often extending over half the length of the abdomen
(Fig.75A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . canatra
- Dorsal cephalic hairs numerous, rather evenly distributed, maximally
70-80 µm long, much longer than the distances between their bases. SIPH
cones higher, with bases much less extensive (Fig. 75B) . . . . . . . . nigra
5 HT I (measured along ventral side) at least 0.5 x HT II (e.g. Fig. 76B).
Longest hairs on dorsal side of hind tibia 110-230 µm long, 1.5 or more
x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- HT I less than 0.5 x HT II. Longest hairs on dorsal side of hind tibia
30-150 µm long, rarely up to 1.5 x middle width of tibia . . . . . . . . . 9
6 ABD TERG 2-7 each with a variably developed but usually large pair of
dark sclerites (or diffuse dusky areas) including the bases of several hairs.
HT II 0.20-0.28 mm. Eyes sessile, always with a distinct ocular tubercle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinivora
- ABD TERG 2-7 with separate small dark hair-bearing sclerites (scleroites).
HT II 0.27-0.53 mm. Eyes on short lateral projections of head, usually
without any evident ocular tubercle (e.g. Fig. 75G) . . . . . . . . . . . . . . . 7
7 Fore tibia uniformly dark, mid- and hind tibia with pale basal sections.
First tarsal segments usually with 2 sense pegs, one behind the other
(Fig. 75H). Largest scleroites on ABD TERG 2-5 of greatest diameter
25-70 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . watsoni
- If the fore tibia is uniformly dark, then so are the mid- and hind tibiae.
First tarsal segments with 1 sense peg. Largest scleroites on ABD TERG
2-5 of greatest diameter 75-300 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 All tibiae with pale section on basal half. First tarsal segments usually with
1-2 dorsal hairs, and with the sense peg some distance from the apex.
ABD TERG 5 with 28-49 hairs between SIPH bases, most of them on
large scleroites (Fig. 80C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
- Tibiae usually rather uniformly dark (sometimes slightly less so near base).
First tarsal segments without dorsal hairs, and with sense peg close to
apex. ABD TERG 5 with 50-110 hairs between SIPH bases, a minority
of which are on large scleroites (Fig. 80D) . . . . . . . . . . . . . . . pergandei
9 Hairs on dorsal side of hind tibia near its midpoint mostly less than 60 µm
in length, rarely up to 70 µm, the longest of them 0.3-0.8 x diameter of
tibia at this point . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Most or all of hairs on dorsal side of hind tibia near its midpoint longer
298 Pinus

Fig. 75. Pattern of dorsal sclerotization of aptera of A, Cinara canatra, B, C. nigra, C, C. brevispinosa
(spring), D, C. contortae, E, C. banksiana, f, C. medispinosa (spring); G, stalked eye of C. pinea;
H, HT I of C. watsoni; hairs on dorsal side of middle part of hind tibia of aptera of I, C. murrayanae
and J, C. atlantica.
 Pinus 299

than 60 µm, the longest of them 0.7-1.5 x diameter of tibia at this point
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
10 ABD TERG 4-6 with many scleroites of width more than 50 µm, or
bars or patches forming part of a more extensive pattern of markings
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- ABD TERG 5 and 6, and usually 4 also, without any bars or patches or
scleroites more than 4 x wider than the hair bases . . . . . . . . . . . . . . 13
11 Length of sclerotized part of stylet groove (Fig. 74A) 1.55-1.76 mm.
Dorsal abdominal markings comprising paired patches of irregular outline
on ABD TERG 1-8, sometimes reduced or broken into groups of sclerites
on 2-6 (Fig. 75D). All tibiae with pale section on basal half . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . contortae
- Length of sclerotized part of stylet groove 1.25-1.54 mm. Dorsal abdomen
either with broad transverse bands on all tergites or with paired patches
on ABD TERG 1-3 and scattered sclerites on 4-5. Fore and hind tibiae
often wholly dark, or with shorter pale section than mid-tibia . . . . . . 12
12 Dorsal abdomen usually (in spring, at least) with a complete pattern of
broad transverse bars on all tergites (Fig. 75C); if reduced on ABD TERG
4-6 then the sclerites are still arranged in single transverse rows across the
midline. Hairs on dorsal side of hind tibia short and spine-like, all less
than 50 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brevispinosa
- Dorsal abdomen with well-developed paired patches on ABD TERG 1 and
2 (or 1-3) and 7; only scattered small sclerites on 4-5, and usually none
at all on 6 (Fig. 75E). Hairs on dorsal side of hind tibia variable, but at
least some are usually more than 50 µm long . . . . . . . . . . . . . banksiana
13 Length of sclerotized part of stylet groove (see Fig. 74A) 1.9 mm or more
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Length of sclerotized part of stylet groove less than 1.8mm . . . . . . 15
14 PT finger-like, parallel-sided, 0.07-0.09 mm long. Length of sclerotized
part of stylet groove 2.3-3.2 mm . . . . . . . . . . . . . . . . . . . . . . . piniradicis
- PT tapering, less than 0.06mm long. Sclerotized part of stylet groove
1.9-2.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schwartzii
15 Length of sclerotized part of stylet groove 1.12-1.37 mm . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ontarioensis
- Length of sclerotized part of stylet groove 1.45-1.65 mm . . . . . taedae
16 Hairs on SIPH cone sparse and very short, the longest about 15 µm long
(only the sexuales of this species are described) . . . . . . . . . parvicornis*
- Hairs on SIPH cone more than 50 µm long . . . . . . . . . . . . . . . . . . . . . 17
17 R IV with 15-28 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
300 Pinus

- R IV with 4-12 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

18 Hind tibia wholly dark and markedly bowed, bearing blunt hairs. R IV
with 15-21 accessory hairs. ABD TERG 3-6 without sclerites. SIPH cones
with less than 40 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . strobi
- Hind tibia with pale section on basal half, and bearing pointed hairs. R
IV with 19-28 accessory hairs. ABD TERG 3-6 usually with sclerites.
SIPH cones with more than 80 hairs . . . . . . . . . . . . . . . . . . . . . cronartii
19 Longest hairs on dorsal side of hind tibia 60-80 µm. Dorsal abdomen
usually with paired dark patches of irregular shape on all tergites
(Fig. 75F; but reduced in midsummer populations, sometimes being
entirely absent from ABD TERG 3-4 and reduced to paired groups of
broken sclerites on 5-6). Length of sclerotized part of stylet groove (see
Fig. 74A) 1.67-1.92 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . medispinosa
- Longest hairs on dorsal side of hind tibia 80-150 µm. Dorsal abdomen
with or without sclerites on ABD TERG 3-6, but if present these are not
in paired groups . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Length of sclerotized part of stylet groove 1.82-2.13 mm . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . oregonensis
- Length of sclerotized part of stylet groove 1.20-1.68 mm . . . . . . . . . 21
21 Hairs on ABD TERG 3-5 maximally 40 µm, each usually with a very small
circular scleroite at its base (Fig. 80G) . . . . . . . . . . . . . . . . . . . . . . . . pini
- Hairs on ABD TERG 3-5 much longer, either not on sclerites or arising
from sclerites or scleroites of varying size and shape . . . . . . . . . . . . . 22
22 ABD TERG 8 usually with a complete transverse band. Longest hairs on
front of head and on ABD TERG 3-5 less than 70 µm long . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . melaina
- ABD TERG 8 usually with paired sclerites not united or only tenuously
united across midline. Longest hairs on front of head and on ABD TERG
3-5 more than 70 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 Hairs on dorsal side of hind tibia maximally 100-150 µm, with finely
pointed apices (Fig. 75I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . murrayanae
- Hairs on dorsal side of hind tibia maximally 80-100 µm, bristle-like, with
blunt or abruptly pointed apices (Fig. 75J) . . . . . . . . . . . . . . . . atlantica

Key to apterae of Essigella spp. on Pinus contorta

1 ABD TERG 2-4 with 6 spinopleural hairs (rarely 7) in a row across the
tergite, and 2 marginal hairs on each side. Dorsal body usually pale, never
very dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . californica
ABD TERG 2-4 with 8-10 (or rarely up to 12) spinopleural hairs, and
3-4 marginal hairs. Body pale or dark . . . . . . . . . . . . . . . . . . . . . . . . . . 2
 Pinus 301

2 Darkest pigmentation of ANT III-V slightly to considerably lighter than

that of ANT I, with ANT III pale except for slight darkening of distal
0.25. Dorsal body, including front of head, uniformly dark brown to
black. Longest hair on dorsal side of hind tibia near midpoint less than
30 µm, usually less than diameter of tibia . . . . . . . . . . . . . . . critchfieldi
Darkest pigmentation of ANT III-V darker than ANT I, with distal
0.3-0.5 of ANT III dark. Dorsal body pale or dark, if dark then front
of head is often paler than dorsal abdomen. Longest hair on dorsal side
of hind tibia near midpoint usually more than 30 µm, usually longer than
diameter of tibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Dorsal body usually moderately to extremely dark, but occasionally pale;
if dark, then frons is usually paler than dorsal abdomen. If pale, the
ventrolateral borders of abdominal tergites, anteroventral border of frons,
and posterior edge of subgenital plate are well defined and demarcated
from adjacent membranous areas. ABD TERG 8 usually with 6 hairs,
rarely up to 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . knowltoni
Dorsal body never very dark, usually pale; if rather dark then front of
head is concolorous with dorsal abdomen. Ventrolateral borders of
abdominal tergites, anteroventral border of frons and/or posterior edge
of subgenital plate often poorly demarcated from adjacent membranous
regions. ABD TERG 8 with 6-8 (rarely up to 10) hairs . . . . . . . braggi

P. cooperi Cinara schwartzii

(Mexican Mountain Pine) Essigella culifornica
(Use key to Cinara spp. on pines of subsect. Ponderosae under P. ponderosa)
P. coulteri Cinara arizonica, diabola,
(Bigcone Pine) montanesa, ponderosae,
tujafilina
Essigella californica, eastopi, fusca
Schizolachnus piniradiatae

Key to Cinara spp. on Pinus coulteri and other pines of subsect. Sabinianae
(sabiniana, torreyana)
1 BL less than 4.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
BL more than 4.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Longest hairs on ANT III 120-200µm, and on hind tibia 140-310 µm
..................................................... tujafilina*
Longest hairs on ANT III 30-60 µm, and on hind tibia 45-80µm . . . . . 3
3 R IV 0.25-0.30 mm. SIPH less than 0.2mm in diameter, bearing a few,
short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . diabola*
R IV 0.17-0.22 mm. SIPH more than 0.5mm in diameter, bearing many
(25 +) long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae
302 Pinus

4 Hairs on dorsal body surface, ANT III-IV and legs all rather sparse,
short, thick and slightly capitate. ANT III longer than IV + V together
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . arizonica
Hairs not capitate; those on hind tibia short (about 50 µm) and dense.
ANT III shorter than IV + V together . . . . . . . . . . . . . . . . . . montanesa

Key to Essigella spp. on Pinus coulteri

1 ABD TERG 2-4 each with 6 spinopleural hairs (rarely 7) in a transverse
row, and 2 marginal hairs on each side . . . . . . . . . . . . . . . . . californica
ABD TERG 2-4 each with 8-12 spinopleural hairs, and 3-5 marginal hairs
on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Dorsal body usually darker than tibiae, with a paler, longitudinal spinal
stripe on thorax and abdomen. Longest hairs on dorsal side of hind tibia
near midpoint 23-83 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . eastopi
Dorsal body pale or dark, if dark then without a paler spinal stripe.
Longest hair on dorsal side of hind tibia near midpoint 50-135 µm . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . fusca

P. cretacea Eulachnus cretacea

P. cubensis Cinara atlantica, pergandei
(Use key to aphids on P. contorta)
P. densiflora Cinara etsuhoe, formosana,
(Japanese Red Pine) orientalls, pergandei,
pinidensiflorae, piniformosana
Eulachnus thunbergii
Pineus havrylenkoi, matsumurae,
orientalis
Prociphilus sp. near laricis
Schizolachnus orientalis
Stomaphis pini

Key to Cinara spp. on Pinus densiflora and other oriental pines of subsect.
Sylvestres (massoniana, tabulaeformis, taiwanensis, thunbergii)
If the outcome of using this key is unsatisfactory, try the key to western
palaearctic species under Pinus sylvestris
1 HT I elongate, 0.16-0.33 mm long (measured on ventral side), at least
0.5 x HT II or longer (Fig. 76B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- HT I less than 0.17 mm long, less than 0.5 x HT II . . . . . . . . . . . . . . 5
2 Hairs on dorsal body and appendages short and thick; longest hairs on
ANT III 55-65 µm, on ABD TERG 3 20-40 µm, on hind tibia 75-85 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . formosana
 Pinus 303

- Hairs much longer; longest on ANT III 90-210 µm, on ABD TERG 3
80-210 µm, on hind tibia 90-230 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Tibiae with pale section on basal half. HT I with 1-2 dorsal hairs
(Fig. 76B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
Tibiae wholly dark, although sometimes slightly less dark near base. HT
I usually without dorsal hairs, rarely with 1 . . . . . . . . . . . . . . . . . . . . . 4
4 ABD TERG 5 with less than 50 hairs between SIPH bases, the longest
80-160 µm. Longest hairs on hind tibia 80-160 µm . . . . . piniformosana
ABD TERG 5 with more than 50 hairs between SIPH bases, the longest
120-200 µm (Fig. 80D). Longest hairs on hind tibia 150-190 µm . . . . . .
...................................................... pergandei
5 R IV 0.44-0.54 mrn long, about as long as ANT III or longer, bearing
28-35 accessory hairs in 4 rows. Longest hairs on dorsal side of hind tibia
shorter than or similar to width oi tibia at midpoint . . . . . . . . . . . . . 6
R IV 0.14-0.22 mm long, less than half as long as ANT III and bearing
7-13 accessory hairs. Longest hairs on dorsal side of hind tibia 1.5 or more
x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6 Hairs on dorsal side of hind tibia small, fine and curved distally so that
they are often almost parallel to tibia; maximally 60-70 µm long, about
0.5 x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . sorini
Hairs on dorsal side of hind tibia fine but erect, the longest 70-80 µm long,
about equal to width of tibia at midpoint . . . . . . . . . . . . . . . . . . etsuhoe
7 ABD TERG 1-2 (or 1-3) with marginal sclerites as well as spinal ones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . orientalis
- ABD TERG 1-3 without marginal sclerites . . . . . . . . . . . . . . . . . . . . . . 8
8 HT II less than 0.3 mm long. ABD TERG 1-7 each with a pair of large
spinal sclerites (sometimes reduced or absent particularly on 2-3). Hind
tibia pale on basal half, contrastingly dark distally . . . . . . . . . . . . . . . . .
................................................. pinidensiflorae
HT II more than 0.3 mm long. ABD TERG 2-6 with only small sclerites.
Hind tibia mainly or wholly dark . . . . . . . . . . . . . . . . . . . . . . californica

P. douglasiana Cinara atlantica, ponderosae

(Douglas Pine) Essigella californica
Eulachnus rileyi
Pineus boerneri
(Use key to Cinara spp. on pines of subsect. Ponderosae under P. ponderosa)
P. durangensis Cinara atlantica
(Durango Pine) Essigella californica
Pineus boerneri
Schizolachnus curvispinosus
304 Pinus

P. echinata Cinara atlantica, melaina,

(Short Leaf Pine) pergandei, watsoni
Eulachnus rileyi
Pineus boerneri
Prociphilus piniradicivora
Schizolachnus piniradiatae

Key to Cinara spp. on Pinus echinata and other pines of subsect. Australes
(caribaea, elliottii, glabra, occidentalis, palustris, pungens, rigida, serotina,
taeda)
1 HT I (measured on ventral side) at least 0.5 x HT II (Fig. 76A, B)
............................................................. 2
- HT I less than 0.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 Longest hairs on dorsal side of hind tibia 70-130 µm, similar to width of
tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maritimae
- Longest hairs on dorsal side of hind tibia 110-230 µm, 1.5 or more x
width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . 4 species (pergandei, pinea, pinivora and watsoni)
thai are separated in couplets 6-8 of the key under P. contorta
3 BL 4.3-6.1 mm. Hairs on dorsal body surface, ANT II1-IV and legs all
short, thick and slightly capitate . . . . . . . . . . . . . . . . . . . . . . . . . arizonica
- BL up to 4.2 mm. Hairs short or long, not capitate . . . . . . . . . . . . . . 4
4 R IV 0.23-0.33 mm long, bearing 19-28 accessory hairs . . . . . . . . . . . 5
- R IV 0.16-0.23 mm long, bearing 6-14 accessory hairs . . . . . . . . . . . . 6
5 ANT VI BASE with 11-16 hairs. Longest hairs on hind tibia 80-110 µm.
Dorsal abdominal hairs long, many of them arising from sclerites of vary-
ing size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cronartii
- ANT VI BASE with 20-23 hairs. Longest hairs on hind tibia 50-75 µm.
Dorsal abdominal hairs short, and at least those on ABD TERG 3-6 not
arising from sclerites (bases sometimes encircled by minute scleroites)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . newelli
6 Subgenital plate with only 2-6 hairs (more in fundatrices). First tarsal
segments with either 2 or 3 sense pegs (ventrally, near apex; see Fig. 75H)
............................................................. 7
- Subgenital plate with 25-50 hairs. First tarsal segments with only 1 sense
peg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 First tarsal segments with 2 sense pegs (as in Fig. 75H) . . . . . . . . westi
- First tarsal segments with 3 sense pegs . . . . . . . . . . . . . . . . . . . . . rigidae
8 ANT VI BASE with 5-8 hairs. R IV with 4-6 accessory hairs. Hind tibia
darker than fore or mid-tibiae. Longest hairs on hind tibia 40-50 µm,
about 0.5 x width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . taedae
 Pinus 305

- ANT VI BASE with 8-16 hairs. R IV with 6-10 accessory hairs. Hind tibia
sometimes with a shorter pale section than fore and mid-tibiae, but not
darker overall. Longest hairs on hind tibia 50-100 µm, 0.75 or more x
width of tibia at midpoint . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 ABD TERG 5 with more than 20 hairs, the longest 60-180 µm, between
SIPH bases (Fig. 80E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
- ABD TERG 5 with less than 20 hairs, maximally 10-80 µm, between SIPH
bases (e.g. Fig. 80F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 Longest hairs on ANT III 25-60 µm, and on dorsal side of hind tibia
50-70 µm. ANT III with 1-3 (non-protruberant) rhinaria near apex
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae
- Longest hairs on ANT III 50-70 µm, and on dorsal side of hind tibia
60-100 µm. ANT III with 0-1 rhinaria near apex (if present, the rhinarium
is small and protuberant) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . melaina

P. edulis Cinara apacheca, atra, caliente,

(Pinyon Pine) edulis, metalica, nitidula,
pinata, pinona, poketa, puerca,
ruslica, tanneri, terminalis,
wahtolca
Essigella hoerneri
Pineus coloradensis
(Use key to Cinara spp. on Pinus subsect. Cembroides under P. cembroides)
P. eldarica Eulachnus tuberculostemmata
P. elliottii Cinara atlanlica, cronartii,
(Longleaf Pitch Pine, Slash Pine) maritimae, melaina, newelli,
pinivora, taedae, watsoni
Essigella californica
Eulachnus rileyi
Pineus boerneri
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
P. engelmanii Essigella californica, fusca
(Engelman's Pine) Pineus boerneri
(Use key to Essigella spp. on pines of subsect. Ponderosae under P. ponderosa)
P. excelsa = P. griffithii
P. flexilis Cinara apini, flexilis, villosa
(Limber Pine) Essigella californica, kirki

Key to Cinara spp. on Pinus flexilis

1 HT II 0.33-0.47 mm long, more than 2 x R IV and densely clothed with
short hairs. R IV with more than 30 accessory hairs . . . . . . . . . flexilis
- HT II 0.24-0.29mm long, less than 1.5 x R IV and normally haired. R
IV with about 10 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
306 Pinus

2 Hairs on hind tibia rather spine-like, those on dorsal side less than 55 µm
long, about 0.5 or less x width of tibia at midpoint . . . . . . . . . . apini
Hairs on hind tibia finer and more numerous, up to 65 µm long, clearly
more than 0.5 x width of tibia at midpoint . . . . . . . . . . . . . . . . . villosa

Key to Essigella spp. on Pinus flexilis

ABD TERG 2-4 each with a transverse row of 6 (rarely 7) spinopleural
hairs, and 2 marginal hairs on each side. ABD TERG 8 with 6-8 hairs
..................................................... californica
ABD TERG 2-4 each with 10-14 spinopleural hairs in two irregular rows,
and 4-6 marginal hairs on each side. ABD TERG 8 with 10-14 hairs
.......................................................... kirki

P. funebris = P. tabulaeformis
P. glabra Cinara arizonica, allantica,
(Spruce Pine) melaina, pergandei, taedae,
watsoni
Eulachnus rileyi
Schizolachnus curvispinosus,
parvus
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
P. gregii Cinara atlantica, maritimae,
ponderosae
(Use key to Cinara spp. on Pinus subsect. Oocarpae under P. oocarpa)
P. griffithii (=P. excelsa = Cinara eastopi, lachnirostris,
P. wallichiana) maculipes, maritimae
(Himalayan Blue Pine) Essigella californica
Eulachnus rileyi
Pineus ghanii, wallichianae
Prociphilus himalayensis
Pseudessigella brachychaeta

Key to Cinara spp. on Pinus griffithii

1 Hairs on at least ABD TERG 3-6 very short, maximally 40 µm or less.
SIPH cones small, with maximum diameter equal to or less than length
of R IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Longest hairs on ABD TERG 3-6 at least 60 µm. Diameter of SIPH cones
much greater than length of R IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 BL 3.7-4.9mm. Legs conspicuously maculate. ANT VI shorter than V
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maculipes
BL 2.4-2.9 mm. Legs not maculate. ANT VI longer than V . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lachnirostris
 Pinus 307

3 BL 4.5-5.2 mm. Dorsal abdomen with dark patches or paired broken

groups of sclerites on ABD TERG 1-8 (Fig. 80A; sometimes reduced on
2-4). Hairs on dorsal side of hind tibia maximally 130-220 µm. R IV with
8-12 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . eastopi
BL 2.5-4.2 mm. Dorsal abdomen with transverse, irregular rows of small
sclerites on ABD TERG 1-6 (Fig. 80B), and transverse bars on 7 and 8.
Hairs on dorsal side of hind tibia maximally 70-130µm. R IV with 4-7
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maritimae

P. halepensis (incl. var. brutia) Cinara balachowskyi, cronartii,

(Aleppo Pine) maghrebica, maritimae,
palaestinensis, pinea, pini
Essigella californica
Eulachnus rileyi,
tuberculostemmata
Pineus boerneri, havrylenkoi

Key to Cinara spp. on Pinus halepensis

1 HT I 0.16-0.33 mm (measured along ventral side), at least 0.5 x HT II
or longer (Figs 76A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- HT I less than 0.14mm long, less than 0.5 x HT II (e.g. Fig. 76C)
............................................................. 3
2 Longest hairs on dorsal side of hind tibia 70-130 µm, shorter than or
about equal to width of tibia at midpoint. HT I 0.16-0.22 mm (Fig. 76A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maritimae
Longest hairs on dorsal side of hind tibia 120-230 µm, often 1.5 x width
of tibia at midpoint or longer. HT I 0.22-0.33 mm (Fig. 76B) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
3 Longest hairs on ABD TERG 3-5 less than 50 µm long . . . . . . . . . . . 4
Longest hairs on ABD TERG 3-5 more than 50 µm long . . . . . . . . . 6
4 R IV 0.39-0.45 mm long, with about 30 accessory hairs . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . balachowskyi
R IV 0.13-0.22 mm long, with 6-10 accessory hairs . . . . . . . . . . . . . . . 5
5 Maximum diameter of SIPH cone 0.09-0.25 mm, 0.3-0.5 x ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . palaestinensis
- Maximum diameter of SIPH cone 0.16-0.70 mm, 0.4-1.4 x ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pini
6 R IV 0.24-0.29 mm long, with 19-28 accessory hairs. Many of hairs on
ABD TERG 2-6 arising from sclerites of various sizes . . . . . cronartii
- R IV 0.13-0.18 mm long, with 6-8 accessory hairs. ABD TERG 2-6
without sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maghrebica
308 Pinus

Fig. 76. Hind tarsus of A, Cinara maritimae, B, C. pinea and C, C. pini.

P. hamata Eulachnus alticola

Pineus pini
P. hartwegii Pineus pini
P. himekomatsu Cinara shinjii
P. insularis Eulachnus thunbergii
P. jeffreyi Cinara arizonica, cronartii,
ponderosae, schwartzii
Essigella californica, fusca,
hillerislambersi
(Use keys to Cinara and Essigella spp. of Pinus subsect. Ponderosae under
P. ponderosa)
P. khasya Aulacorthum (Neomyzus)
(Khasya Pine, Philippine Pine) circumflexum
Cinara atrotibialis, pinea
Eulachnus thunbergii
Pineus pini
Key to Cinara spp. on Pinus khasya
Fore tibia wholly black, mid- and hind tibiae also black or with slightly
paler section on basal half. R IV 0.26-0.38 mm long, at least 0.5 x ANT
III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atrotibialis
Fore tibia often darker than mid- and hind tibiae, but all tibiae with paler
section on basal half. R IV 0.21-0.29 mm long, less than 0.5 x ANT III
.......................................................... pinea
 Pinus 309

P. koraiensis Cinara cembrae,

(Korean Pine) minoripinihabitans, ?mongolica,
[pinikoraiensis], [taeniatoides
Mordvilko, 1901], watanabei
Eulachnus pumilae
Pineus cembrae, cembrae ssp.
pinikoreanus, cladogenous,
cortecicolus, hosoyai
(Use key to Cinara spp. on Pinus subsect. Cembrae under P. cembra)
P. lambertiana Cinara hirticula, moketa,
(Sugar Pine) saccharinipini
Essigella californica, kathleeni
Pineus coloradensis
Prociphilus caryae ?ssp.
arbutifoliae

Key to Cinara spp. on Pinus lambertiana

Of the three species described by Hottes (1957a, 1958a) from P. lambertiana,
the available specimens all seem to correspond most closely to moketa; couplet
2 is based solely on the original descriptions.
1 Longest hairs on ANT III 100-110µm, on dorsal side of hind tibia
100-120 µm, and on ABD TERG 5 and SIPH cones 120-150 µm . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . moketa
Longest hairs on ANT I I I 80-90µm, on dorsal side of hind tibia
70-100 m, and on ABD TERG 5 and SIPH cones 90-130 µm . . . . . 2
2 Longest hairs on dorsal side of hind tibia about 100µm. HT I about
0.5 x HT II. HT II about 1.3 x R IV. R IV about 0.6 x maximum
diameter of SIPH cone . . . . . . . . . . . . . . . . . . . . . . . . . . . . saccharinipini *
Longest hairs on dorsal side of hind tibia about 70 µm. HT I about
0.4 x HT II. HT II about 1.8 x R IV. R IV about 0.3 x maximum
diameter of SIPH cone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hirticula*

Key to Essigella spp. on Pinus lambertiana

ABD TERG 2-4 each with a transverse row of 6 spinopleural hairs (rarely
7), and with 2 marginal hairs on each side. ABD TERG 8 with 6-8 hairs.
HT II less than 1.9 x HT I . . . . . . . . . . . . . . . . . . . . . . . . . . . . californica
ABD TERG 2-4 each with 11-14 spinopleural hairs, in two irregular rows,
and 4-5 marginal hairs on each side. ABD TERG 8 with 7-13 hairs. HT
II more than 1.9 x HT I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . kathleenae

P. laricio = P. nigra maritima

P. leiophylla Cinara terminalis
310 Pinus

Essigella californica, fusca

Pineus pini group
(For separation of the two Essigella species use the key under P. coulteri)
P. leucodermis Cinara schimitscheki
(or try key to Cinara on pines of subsect. Sylvestres under P. sylvestris)
P. luchuensis Eulachnus thunbergii
P. lumholtzii Cinara atlantica
P. lutea Pineus pini group
P. massoniana Cinara pinidensiflorae
(Chinese Red Pine) Eulachnus rileyi, thunbergii
Pineus pini group
Schizolachnus orientalis
(Use key to Cinara spp. on P. densiflora)
P. maximinoi Pineus pini group
P. merkusii Eulachnus thunbergii
Pineus pini group
P. michoacana Cinara ponderosae, terminalis
Essigella californica
Eulachnus rileyi
Pineus boerneri
(Use key to Cinara spp. on Pinus subsect. Ponderosae under P. ponderosa)
P. monophylla Cinara atra, caliente, edulis,
(Single-leaf Pinyon Pine) tanneri, wahtolca
Essigella hoerneri
Pineus coloradensis
(Use key to Cinara spp. on Pinus subsect. Cembroides under P. cembroides)
P. montezumae Pineus pini group
P. monticola Cinara ferrisi, hirsuta, kuchea
(Western White Pine) Essigella californica, kathleeni
Pineus coloradensis, pinifoliae
Prociphilus caryae ssp. caryae
(For separation of Essigella spp. see key under P. lambertiana)

Key to Cinara spp. on Pinus monticola

1 BL more than 4.5 mm. Tibiae wholly black. Length of sclerotized part of
stylet groove (see Fig. 74A) more than 2.4mm . . . . . . . . . . . . . . . ferrisi
BL less than 4.5 mm. Tibiae with paler section on basal half. Length of
sclerotized part of stylet groove less than 2.0 mm . . . . . . . . . . . . . . . . . 2
2 Longest hairs on dorsal side of hind tibia 70-80 µm long . . . . . . . kuchea
 Pinus 311

Longest hairs on dorsal side of hind tibia up to 150 µm long . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hirsuta

P. mugo (incl. var. mughus, var. Cinara covassii, guadarramae,

pumilio) hyperophila, neubergi,
(Mountain Pine) pergandei, pinea, pini,
pinihabitans, setosa
Eulachnus alticola, brevipilosus,
intermedius, rileyi
Pineus orientalis, pini
Prociphilus (Stagona) pini
(Use key to Cinara spp. on Pinus subsect. Sylvestres under P. sylvestris)
P. muricata Cinara atlantica
(Bishop Pine) Essigella californica
Pineus pini group
(Use key to Cinara spp. on Pinus subsect. Oocarpae under P. oocarpa)
P. nigra (incl. var. caramatica, var. Cinara acutirostris, atlantica,
maritima) brauni, laricionis, maritimae,
(Austrian Pine) [montanensis], pinea, pini,
ponderosae, schimilscheki
Eulachnus agilis, brevipilosus,
mediterraneus, nigricola, rileyi
Pineus coloradensis, pini group
Schizolachnus obscurus, pineti
(Use key to Cinara spp. on Pinus subsect. Sylvestres, under P. sylvestris)
P. occidentalis Cinara arizonica, atlantica,
(West Indian Pine) ponderosae
Essigella californica
(Use key to Cinara spp. on Pinus subsect, Australes under P. echinata)
P. oocarpae Cinara atlantica, azteca
(Ocote Pine) Pineus boerneri

Key to Cinara spp. on Pinus oocarpae and other pines of subsect. Oocarpae
(attenuata, greggii, muricata, patula, radiata)
1 R IV 0.23-0.33 mm long, bearing 14-28 accessory hairs . . . . . . . . . . . 2
R IV 0.14-0.23 mm long, bearing 6-10 accessory hairs . . . . . . . . . . . . 3
2 Longest hairs on ANT III 60-80 µm, on hind tibia 50-80 µm . (ABD TERG
2-6 without sclerites?) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . essigi*
Longest hairs on ANT III and hind tibia 80-110 µm, all very fine. (ABD
TERG 2-6 often with numerous small sclerites) . . . . . . . . . . . . cronartii
3 HT I 0.16-0.22 mm long, more than 0.5 x HT II (Fig. 76A) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maritimae
312 Pinus

- HT I less than 0.14mm long, less than 0.5 x HT II . . . . . . . . . . . . . . 4

4 Maximum diameter of SIPH cone 0.3-0.4 mm. Hairs on ABD TERG 5
about 70 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . azteca*
Maximum diameter of SIPH cone usually in range 0.4-0.7 mm. Hairs on
ABD TERG mostly either shorter or longer than 70 µm . . . . . . . . . . . 5
5 SIPH with more than 60 hairs, including many on basal part of cone.
ABD TERG 5 with more than 25 hairs, 60-110 µm long, between SIPH
cones (Fig. 80E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
SIPH with less than 60 hairs, rather sparse on basal part of cone. ABD
TERG 5 with 4-15 hairs, 10-60 µm long, between SIPH cones (Fig. 80F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae/terminalis

P. palustris Cinara atlantica, melaina, watsoni

(American Pitch Pine, Longleaf Essigella californica
Pine) Eulachnus rileyi
(Use key to Cinara spp. on Pinus subsect. Australes, under P. echinata)
P. parviflora Cinara shinjii, watanabei
(Japanese White Pine) Eulachnus pumilae
Pineus cembrae, harukawai,
havrylenkoi

Key to Cinara spp. on Pinus parviflora

R IV more than 0.25mm long, bearing 14-22 accessory hairs. Hairs on
ABD TERG 5 less than 70 µm long. ABD TERG 2-5 with only small
scleroites ringing hair bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . watanabei
R IV less than 0.25 mm long, bearing 6-8 accessory hairs. Hairs on ABD
TERG 5 up to 120 µm long. ABD TERG 2-5 with large sclerites . . . . . . .
......................................................... shinjii

P. patula Cinara atlantica, cronartii,

(Mexican Weeping Pine) terminalis
Essigella californica
Eulachnus rileyi
Pineus boerneri
(Use key to Cinara spp. on pines of subsect. Oocarpae under P. oocarpa)
P. pentaphylla = P. parviflora
P. peuce Cinara cembrae
Pineus strobi
P. pinaster Cinara guadarramae, maghrebica,
(Maritime Pine, Cluster Pine) maritimae
Eulachnus agilis, brevipilosus,
 Pinus 313

mediterraneus, rileyi,
tuberculostemmata
Pineus harukawai
(Use key to Cinara spp. on western palaearctic pines of subsect. Sylvestres,
under P. sylvestris)
P. pinea Cinara acutirostris, maghrebica,
(Italian Stone Pine) maritimae, schimitscheki
Essigella californica
Eulachnus rileyi,
tuberculostemmata
Pineus boerneri, havrylenkoi
Schizolachnus obscurus
(The three Cinara species are separated in the key under P. sylvestris)
P. ponderosa (incl. var. arizonica, Cinara arizonica, glabra,
var. scopularum) nigrita(?), oregonensis, pinea,
(Ponderosa Pine) pini, ponderosae,
pseudoschwartzii, schwartzii,
solitaria, thatcheri
Essigella californica, essigi, fusca
Eulachnus rileyi
Pineus boerneri, coloradensis
Schizolachnus curvispinosus,
flocculosus, piniradiatae

Key to Cinara spp. on Pinus ponderosa and other pines of subsect. Ponderosae
(cooperi, douglasiana, durangensis, jeffreyi, michoacana, pseudostrobus)
1 BL 4.6-6.Omm. Hairs on dorsal body surface, ANT III-IV and legs all
rather widely spaced, short, thick and slightly capitate . . . . . arizonica
- BL less than 4.6mm. Hairs sometimes short and thick but not widely
spaced and not capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 HT I 0.15-0.23 mm long, more than 0.5 x HT II (Fig. 76B). Hairs on hind
tibia 120-230 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
- HT I 0.08-0.15 mm long, less than 0.5 x HT II. Hairs on hind tibia
20-120 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 R IV with 12-28 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- R IV with 6-10 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4 Longest hairs on hind tibia 30-60 µm, about 0.5 x width of tibia at mid-
point or shorter. Hairs on ABD TERG 5 mostly short and blunt, less than
30 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Longest hairs on hind tibia 70-120 µm, clearly more than 0.5 x width of
tibia at midpoint. Hairs on ABD TERG 5 with acute apices, the longest
at least 70 µm in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
314 Pinus

Fig. 77. Dorsal abdomen of aptera of Cinara glabra.

5 Dorsum with a large dark sclerite fused across midline and extending
across ABD TERG 2-1, but not fused with SIPH bases (Fig. 77). ABD
TERG 5 with more than 50 hairs between SIPH bases. Ventral surface
covered with dense rows of pointed spicules . . . . . . . . . . . . . . . . . glabra
- Dorsum only lightly sclerotized, often with dusky areas on each side of
midline. ABD TERG 5 with less than 20 hairs between SIPH bases. Ven-
tral surface only faintly and minutely spiculose . . . . . . . . . . . . . . . . . . 6
6 R IV 0.20-0.25 mm long, with 15-16 accessory hairs. ANT V about equal
in length to ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schwartzii
- R IV 0.26-0.30 mm long, with 19-21 accessory hairs. ANT V clearly
longer than ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . thalcheri
7 ABD TERG 2-6 without distinct sclerites but often with a dusky area on
each side of midline. ABD TERG 5 with less than 20 hairs between SIPH
bases. R IV 0.22-0.24 mm long . . . . . . . . . . . . . . . . . . . pseudoschwartzii
- ABD TERG 2-6 with scattered separate, distinct sclerites. ABD TERG 5
with more than 50 hairs between SIPH bases. R IV 0.24-0.29 mm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cronartii
8 SIPH cones with two distinctly different types of hair, one type long
(about 70 µm) and spine-like, the other small (about 12 µm), fine and
numerous. R IV 0.22-0.26 mm long. Body rounded, tick-like, tibiae short
and black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . solitaria
- SIPH cones with one type of hair. R IV 0.14-0.22. Body oval . . . . 9
 Pinus 315

9 Longest hairs on hind tibia 80-110 µm, and on ABD TERG 5, 60-110µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Longest hairs on hind tibia 40-95 µm, and on ABD TERG 5, 10-70 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Length of sclerotized part of stylet groove (see Fig. 74A) 1.8-2.2 mm.
ABD TERG 5 with less than 20 hairs between the rather small SIPH cones,
each bearing 15-20 hairs. ABD TERG 7 and 8 with broad transverse bars
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . oregonensis
- Length of sclerotized part of stylet groove 1.3-1.7 mm. ABD TERG 5 with
more than 25 hairs (Fig. 80E), between large SIPH cones each bearing
more than 50 hairs. ABD TERG 7 and 8 with paired sclerites not joined
or only tenuously linked across midline . . . . . . . . . . . . . . . . . . . atlantica
11 Length of sclerotized part of stylet groove (see Fig. 74A) 1.5-2.0mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae/terminalis
- Length of sclerotized part of stylet groove 1.2-1.4 mm . . . . . . . . . pini

Key to Essigella spp. on pines of subsect. Ponderosae

1 ABD TERG 2-4 each with a transverse row of 6 (rarely 7) spinopleural
hairs, and 2 marginal hairs on each side . . . . . . . . . . . . . . . . californica

ABD TERG 2-4 each with 8-12 spinopleural hairs (rarely 7), and 3-5
(rarely 2) marginal hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2 Metathoracic tergite fused with ABD TERG 1, either entirely across dor-
sum (pale individuals) or at least laterally (dark individuals). All tibiae
dusky to dark, concolorous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . essigi
Metathoracic tergite not fused to ABD TERG 1, even laterally. Fore and
hind tibiae clearly darker than middle tibia . . . . . . . . . . . . . . . . . . . . . . 3
3 BL 2.10-2.64 mm. ANT III 0.16-0.23 mm. Hind tibia 1.06-1.56 mm.
Dorsal body pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hillerislambersi
- BL 1.79-2.39mm. ANT III 0.12-0.18mm. Hind tibia 0.76-1.13mm.
Dorsal body pale or dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . fusca

P. pseudostrobus Cinara terminalis

(False Weymouth Pine) Essigella californica
Pineus boerneri
(Use key to Cinara spp. of Pinus subsect. Ponderosae under P. ponderosa)
P. pumila Cinara ?mongolica
(Japanese Stone Pine) Eulachnus pumilae, thunbergii
Pineus cembrae
(Use key to Cinara spp. on Pinus subsect. Cembrae under P. cembra]
316 Pinus

P. pungens Cinara atlantica, pinivora, taedae,

(Hickory Pine, Table Mountain watsoni
Pine) Pineus boerneri
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
P. quadrifolia Cinara ponderosae
(California Pinyon Pine) Essigella hoerneri
(Use key to Cinara spp. on Pinus subsect. Cembroides under P. cembroides)
P. radiata Cinara atlantica, maritimae
(Monterey Pine, Insignis Pine) Essigella californica, essigi
Eulachnus rileyi
Pineus boerneri, pini, sylvestris
Schizolachnus pineti, piniradiatae
(Use key to Cinara spp. on Pinus subsect. Oocarpae under P. oocarpa)
(The two Essigella species are separated in the key under P. ponderosa)
P. resinosa Cinara atlantica, harmonia,
(Canadian Red Pine) melaina, pinea, pinivora, slrobi
Pineus boerneri, coloradensis
Schizolachnus piniradiatae

Key to Cinara spp. on Pinus resinosa

1 HT I (measured along ventral side) more than 0.5 x HT II (e.g. Fig. 76B).
Longest hairs on dorsal side of hind tibia 110-230 µm long . . . . . . . . 2
HT I about 0.5 x HT I I or less. Longest hairs on dorsal side of hind tibia
60-130 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 HT I 0.12-0.15 mm, HT II 0.20-0.28 mm. ABD TERG 2-7 each with a
variably developed but usually large pair of sclerites (or diffuse dusky
areas) including the bases of several hairs . . . . . . . . . . . . . . . . . pinivora
- HT I 0.17-0.33 mm, HT II 0.35-0.53 mm. ABD TERG 2-7 with separate
small hair-bearing scleroites (Fig. 80C) . . . . . . . . . . . . . . . . . . . . . . . pinea
3 R IV bearing 15-22 accessory hairs. Hind tibia wholly dark and often
markedly curved, bearing blunt hairs . . . . . . . . . . . . . . . . . . . . . . . strobi
R IV bearing 6-10 accessory hairs. Hind tibia with a pale section on basal
half and bearing pointed hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 ABD TERG 8 usually with a continuous transverse dark bar. Hairs on
front of head maximally 70 µm, mostly shorter than or about equal to
width of ANT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . melaina
ABD TERG 8 usually with transverse bar interrupted in or only tenuously
linked across midline. Hairs on front of head maximally 80-120µm,
mostly longer than width of ANT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 ABD TERG 8 with 12-16 hairs. BL 2.2-3.3 mm, 4.5-5.6 x maximum
width of SIPH cone. Longest hairs on dorsal side of hind tibia 80-100 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
 Pinus 317

- ABD TERG 8 with 17-23 hairs. BL 2.9-4.2mm, 5.7-9.0 x maximum

width of SIPH cone. Longest hairs on dorsal side of hind tibia 90-130 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . harmonia

P. rigida Cinara atlantica, pergandei, pinea,

(Pitch Pine) [pinidensiflorae], pinivora,
rigidae, taedae, watsoni
Eulachnus thunbergii
Pineus boerneri, coloradensis
Prociphilus piniradicivorus
(Use key to Cinara spp. of Pinus subsect. Australes under P. echinata)
P. roxburghii Cinara atlantica, atrotibialis,
(Asiatic Longleaf Pine, Cher Pine) ponderosae
Essigella californica
Pineus simmondsi
Schizolachnus curvispinosus

Key to Cinara spp. on Pinus roxburghii

1 Tibiae wholly dark, although sometimes with slightly less dark section on
basal half of mid- and hind tibiae. HT I 0.20-0.24 mm, more than
0.5 x HT II. R IV 0.26-0.38 mm long . . . . . . . . . . . . . . . . . . atrotibialis
All tibiae with pale section on basal half. HT I 0.08-0.12 mm long, less
than 0.5 x HT II. R IV 0.16-0.21 mm long . . . . . . . . . . . . . . . . . . . . . . 2
2 ABD TERG 5 with more than 20 hairs between SIPH bases, the longest
60-80 µm (Fig. 80E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
- ABD TERG 5 with less than 20 hairs between SIPH bases, the longest
10-60 µm long (Fig. 80F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae

P. rudis Pineus boerneri

P. sabiniana Cinara arizonica, ponderosae
(Digger Pine) Essigella californica
Pineus boerneri
Schizolachnus piniradiatae
(Use key to Cinara spp. of Pinus subsect. Sabinianae under P. coulteri)
P. serotina Cinara atlantica, cronartii, taedae,
(Pond Pine) watsoni
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
P. sibirica Cinara mongolica
(Siberian Stone Pine) Pineus cembrae
(Use key to Cinara spp. on Pinus subsect. Cembrae under P. cembra)
P. strobiformis Cinara flexilis
Essigella californica, kirki
(Use keys to Cinara and Essigella spp. on P. flexilis)
318 Pinus

P. strobus Cinara atlantica, pinea, strobi

(Eastern White Pine, Weymouth Essigella californica, pini
Pine) Eulachnus pumilae, rileyi
Pineus boerneri, coloradensis,
floccus, harukawai, pini,
pinifoliae, strobi
Prociphilus caryae ssp. fitchii
Prociphilus (Stagona)
crataegistrobi
(For separation of the three Cinara species use the key under P. resinosa)

Key to Essigella spp. on Pinus strobus

CAUDA with a moderately to well-developed apical tubercle, usually as
long as its basal width or longer. Longest hairs on dorsal side of hind tibia
near its midpoint less than 30 µm long. R IV+V 0.055-0.075 mm long . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pini
CAUDA with apical tubercle weakly developed (not as long as its basal
width). Longest hairs on dorsal side of hind tibia near midpoint variable
but usually longer than 30 µm. R IV+V 0.070-0.098 mm long . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . californica

P. sylvestris Cinara atlantica, formosana,

(Scots Pine) guadarramae, hyperophila,
intermedia, [longirostris], nuda,
[oblonga (del Guercio, 1909)],
pilosa, pinea, pini,
piniformosana, pinihabitans,
piniphila, pinivora, taeniata,
watanabei
Eulachnus agilis, brevipilosus,
rileyi, thunbergii
Pineus boerneri, coloradensis,
orientalis, pini, strobi, sylvestris
Prociphilus (Stagona) pini
Schizolachnus pineti

Key to Cinara spp. on Pinus sylvestris and other western palaearctic pines of
subsect. Sylvestres (mugo, nigra, pinaster); also including species on P. pinea.
(Cinara spp. on P. halepensis, including var. brutia, are keyed separately
under that host plant.)
1 ABD TERG 5-7 usually with an extensive dark sclerite encompassing
SIPH cones (e.g. Fig. 78; the lateral extent of the sclerite varies con-
siderably, as does the occurrence of clear areas within it) . . . . . brauni
- ABD TERG 5-7 with or without sclerites but these are never fused
between segments or with SIPH cones . . . . . . . . . . . . . . . . . . . . . . . . . . 2
 Pinus 319

Fig. 78. Dorsal abdomen of aptera of Cinara brauni.

2 HT I (measured along ventral side) 0.5 or more x HT II (e.g. Figs76A,

B); but if almost exactly 0.5 X then HT I is more than 0.16mm long
............................................................. 3
- HT 1 0.07-0.17 mm long and less than 0.5 x HT II (e.g. Fig. 76C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3 HT I 0.12-0.15 mm, HT II 0.20-0.28 mm . . . . . . . . . . . . . . . . . . . . . . . 4
- HT I 0.16-0.32 mm, HT II 0.30-0.53 mm . . . . . . . . . . . . . . . . . . . . . . . 5
4 Rostrum much shorter than body. ABD TERG 1-7 each with a pair of
variably developed but usually large sclerites (often irregular in outline or
fragmented) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinivora
- Rostrum long, extending beyond end of abdomen. ABD TERG 2-6
without large sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedia
5 R IV 0.29-0.36mm long, 0.9 or more x HT II. Hairs on ABD TERG
1-7 (Fig. 79A) all arising from small dark circular scleroites of very
uniform size (20-25 µm in diameter) . . . . . . . . . . . . . . . . . . schimitscheki
- R IV 0.15-0.32 mm long, 0.8 or less x HT II. Hairs on ABD TERG 1-7
not all arising from circular, uniform-sized scleroites of diameter
20-25 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Hairs on ABD TERG 5 maximally 27-42 µm, all with very small basal
scleroites (diameter less than 10 µm) . . . . . . . . . . . . . . . . . . . . . . . covassii
- Hairs on ABD TERG 5 maximally 70-230 µm, some of them arising from
much larger scleroites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Longest hairs on ABD TERG 3-5 70-80 µm long, the lengths of the
longest hairs on the largest scleroites mostly not exceeding the maximum
320 Pinus

Fig. 79. Hair-bearing scleroites on ABD TERG 5 of aptera of A, Cinara schimitscheki and B, C. piniphila.

diameter of those scleroites (Fig. 79B). Longest hairs on ANT III

60-70 µm, on dorsal side of hind tibia 60-90 µm . . . . . . . . . . . piniphila
- Longest hairs on ABD TERG 3-5 80-230 µm long, those on the largest
scleroites usually greatly exceeding the diameter of the scleroites (except
when the scleroites join two or more hair bases). Longest hairs on ANT
III 60-210 µm, on dorsal side of hind tibia 70-230 µm . . . . . . . . . . . . 8
8 Largest sclerites on ABD TERG 2-4 of maximum diameter 70-300 µm (if
only 70 µm then there are several of this diameter). ANT II bearing 5-10
hairs and ANT BASE VI with 2-8 hairs . . . . . . . . . . . . . . . . . . . . . . . . 9
- Largest sclerites on ABD TERG 2-4 of maximum diameter 25-70 µm, but
only rarely exceeding 60 µm. ANT II bearing 7-14 hairs and ANT BASE
VI bearing 6-12 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
9 R IV 1.2-1.5 x HT I. HT II 1.3-1.6 x R IV . . . . . . . . . . . . . . neubergi
- R IV 0.7-1.2 x HT I. HT II 1.4-2.2 x R IV . . . . . . . . . . . . . . . . . . . 10
10 Tibiae rather uniformly dark (sometimes less so near bases). First tarsal
segments without dorsal hairs. ABD TERG 5 with 50-110 hairs between
SIPH cones, a minority of which are on large scleroites (Fig. 80D)
...................................................... pergandei
- Tibiae usually at least with pale section on basal half. First tarsal segments
with 1-7 dorsal and dorsolateral hairs. ABD TERG 5 with 25-49 hairs
between SIPH cones, most of them on large scleroites (e.g. Fig. 80C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
 Pinus 321

11 First tarsal segments with 4-7 dorsal and dorsolateral hairs. HT I

0.23-0.32 mm long. ANT V usually without a secondary rhinarium (rarely
1 on one antenna) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pilosa
- First tarsal segments with 1-3 dorsal hairs (Fig. 76B). HT I 0.18-0.26 mm
long. ANT V usually with 1-2 secondary rhinaria on each antenna, rarely
with 0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinea
12 HT II 1.7-2.4 x R IV, which is 0.15-0.21 mm long. Tibiae black . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . piniformosanus
- HT II 1.1-1.6 x R IV, which is 0.25-0.30 mm long. Tibiae with pale basal
section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 ABD TERG 5 with 10-27 hairs between SIPH cones (more in fundatrices).
Length of antennal flagellum (ANT III-VI inclusive) 0.45-0.70 x BL. HT
I 0.19-0.25 mm. Longest hairs on dorsal side of hind tibia 90-180 µm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . laricionis
- ABD TERG 5 with 30-60 hairs between SIPH cones (Fig. 80B). Length
of antennal flagellum 0.34-0.50 x BL. HT I 0.16-0.22 mm. Longest hairs
on dorsal side of hind tibia 60-130µm . . . . . . . . . . . . . . . . . . maritimae
14 R IV more than 0.25mm long, bearing 14-22 accessory hairs . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . watanabei
- R IV less than 0.25 mm long, bearing 4-12 accessory hairs . . . . . . . 15
15 Longest hairs on ANT III 110-150µm, and on hind tibia 170-190µm.
R IV with 4-6 accessory hairs. ABD TERG 5 with 30-50 hairs, the longest
140-180 µm, between SIPH cones . . . . . . . . . . . . . . . . . . . . . pinihabitans
- Longest hairs on ANT III 25-120 µm, on hind tibia 40-130 µm. R IV with
4-12 accessory hairs. ABD TERG 5 with 3-35 hairs between SIPH cones,
the longest 5-140 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 R IV bearing 4-6 accessory hairs, and less than 0.6 x HT II, which is
0.31-0.38 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hyperophila
- R IV bearing 6-12 hairs, and more than 0.6 x HT II, which is
0.19-0.36 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 R IV+V 0.29-0.38 mm, 1.2-1.5 x HT II. ABD TERG 5 with 4-7 hairs
(but up to 18 in fundatrices), 40-110 µm long, between SIPH cones . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . acutirostris
- R IV+V 0.20-0.34mm, 0.9-1.2 x HT II. ABD TERG 5 with 3-35 hairs
between SIPH cones, but if less than 10 then they are usually shorter than
40 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 ABD TERG 2-6 with at least some hairs arising from dark, irregular-
shaped sclerites of diameter 5 or more times that of the hair bases. ABD
TERG 7 as well as 8 usually with dark transverse bars of paired dark
patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
322 Pinus

- ABD TERG 2-6 without irregular-shaped sclerites, although often with

small circles of pigment around hair bases. ABD TERG 7 with or without
large sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19 SIPH with more than 60 hairs, including many on basal part of cone.
ABD TERG 5 with more than 25 hairs, 60-110 µm long, between SIPH
cones (Fig. 80E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . atlantica
- SIPH with less than 60 hairs, which are rather sparse on basal part of
cone. ABD TERG 5 with 4-15 hairs, 10-60 µm long, between SIPH cones
(Fig.80F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ponderosae
20 Sclerotized part of stylet groove (see Fig. 74A) 1.8-2.2 mm long. ANT PT
with 4-7 subapical hairs. ABD TERG 5 with 5-6 hairs (up to 18 in funda-
trices), which are very short and thin, the longest 13-30µm . . . . . . nuda
- Sclerotized part of stylet groove 1.0-1.7 mm long. ANT PT with only 4
subapical hairs. ABD TERG 5 with 3-17 hairs (more in fundatrices), the
longest 15-130µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Length of Sclerotized part of stylet groove 1.0-1.2 mm Diameter of SIPH
cone 0.12-0.40 mm. ABD TERG 5 with 12-18 long, fine hairs (more in
fundatrices), the longest 60-130 µm, between SIPH cones. Longest hairs
on hind tibia 60-130 µm. ABD TERG 7 as well as 8 usually with a pair
of dark patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maghrebica
- Length of Sclerotized part of stylet groove 1.2-1.7 mm. Diameter of SIPH
cone 0.24-0.70 mm. ABD TERG 5 with 3-11 hairs (up to 30 in fundatrices),
the longest 15-100µm, between SIPH cones. Longest hairs on hind tibia
40-95 µm. ABD TERG 7 usually without any large sclerites . . . . . . . . 22
22 ABD TERG 5 with 3-8 hairs (up to 30 in fundatrices), the longest
15-35 µm, between SIPH cones (Fig. 80G). Length of sclerotized part of
stylet groove 1.2-1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pini
- ABD TERG 5 with 6-11 hairs, the longest 35-100 µm, between SIPH
bases (Fig. 80H). Length of sclerotized part of stylet groove 1.3-1.75 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . guadarramae

P. tabulaeformis Cinara californica, formosana,

(Chinese Pine) pinea, pinidensiflorae,
piniformosana
Eulachnus sp.
Pineus boerneri
(Use key to Cinara spp. on oriental pines of subsect. Sylvestres under
P. densiflora)
P. taeda Cinara atlantica, cronartii,
(Loblolly Pine, Carolina Pine) gracilis, melaina, newelli,
pergandei, pinea, pinivora,
taedae, watsoni, westi
Essigella californica, pini
Eulachnus rileyi, thunbergii
 Pinus 323

Fig. 80. Region of ABD TERG 5 between SIPH of aptera of A, Cinara eastopi, B, C. maritimae,
C, C. pinea, D, C. pergandei, E, C. atlantica, f, C. ponderosae, 6, C. pini, H, C. guadarramae.

Pineus boerneri
Prociphilus carolinensis,
piniradicivorus
Schizolachnus parvus
(Use key to Cinara spp. on Pinus subsect. Australes under P. echinata)
(To separate the two Essigella species use the key under P. strobus)
324 Pinus

P. taiwanensis Eulachnus rileyi, thunbergii

(If Cinara spp. are found try key under P. densiflora)
P. tenuifolia Benth. nec Salisb. = P. maximinoi
P. teocote Pineus boerneri
P. thunbergii Cinara etsuhoe, formosana,
(Japanese Black Pine) pinidensiflorae, piniformosana,
sorini
Essigella californica
Eulachnus rileyi, thunbergii
Pineus laevis, orientalis, pini,
sylvestris
Prociphilus sp. near laricis
(Use key to Cinara spp. on oriental pines of subsect. Sylvestres under
P. densiflorá)
P. torreyana Essigella californica
P. tuberculata = P. radiata
P. virginiana Cinara atlantica, gracilis, melaina,
(Virginia Pine) pergandei, pinea, pinivora,
schwartzii, taedae, watsoni
Essigella pini
Eulachnus rileyi
Pineus boerneri
Schizolachnus lanosus
(Use key to Cinara spp. on Pinus subsect. Contortae under P. contorta)
P. wallichiana = P. griffithii
P. yunnanensis Cinara formosana, orientalis ssp.
lijiangensis
(Use key to Cinara spp. on oriental pines of subsect. Sylvestres under
P. densiflora)
Pinus spp.
The following aphids were described from unidentified species of Pinus.
Cinara brevipilosa, orientalis,
paxilla, saraswatae, subapicula,
tenuipes, tibetapini
Pineus pinicorticis
[Prociphilus pini Tao, 1970]
 Pistacia 325

PIPTOCARPHA Compositae

Piptocarpha rotundifolia Aphis gossypii

PISONIA Nyctaginaceae

Pisonia fragrans Toxoptera aurantii

Pisonia sechellarum Toxoptera aurantii

PISTACIA Anacardiaceae

Pistacia spp. are the primary hosts of Fordina (Pemphiginae), which produce
a wide variety of galls, sometimes very large and elaborate (Fig. 126). No other
aphids (apart from the polyphagous Aphis gossypii) are recorded from this
genus. Following the host plant list, two keys are provided, both taking the
genus Pistacia as a whole: the first key is to the emigrant alatae produced in
the galls, which usually open in autumn, and the second is to the alate
sexuparae, which may be collected on the trunks when they return to Pistacia
in spring. Confirmation of identity will often be possible from the description
of the gall given in the systematic section of the book; many of the galls are
illustrated in Fig. 126.

Host Plant List

Pistacia atlantica [Aphis gossypii]
(Betoum) Forda hirsuta, riccobonii
Fordini 'sp. B' of Koach and
Wool (1977)
Geoica harpazi, mimeuri, rungsi
Slavum wertheimae
Smynthurodes betae
P. bohemica ? Geoica lucifuga
P. ethiopica (?) Aloephagous myersi
P. fastula Baizongia pistaciae
326 Pistacia

P. integerrima Baizongia pistaciae

Forda sp(p).
P. khinjuk Asiphonella cynodonti
Baizongia pistaciae
Forda formicaria, kaussarii,
sp. near sichangensis
Geoica lucifuga, setulosa
Paracletus cimiciformis
Rectinasus buxtoni
P. lentiscus Aploneura lentisci
Forda marginata
[Geoica utricularia]
P. mexicana Geopemphigus sp.
P. mutica Forda formicaria, hirsuta,
?riccobonii
Geoica muticae
Slavum esfandiarii, lentiscoides,
wertheimae
Smynthurodes betae
P. palaestina [Aphis gossypii]
Asiphonella cynodonti
Baizongia pistaciae
Forda formicaria, marginata
Geoica sp. nr utricularia
Paracletus cimiciformis
Rectinasus buxtoni
P. sinensis Asiphonella dactylonii
Baizongia pistaciae
Chaetogeoica foliodentata
Forda sichangensis
Geoica lucifuga
P. terebinthus Baizongia pistaciae
Forda formicaria, hirsuta,
marginata
Geoica mimeuri, utricularia
Paracletus cimiciformis,
donisthorpei
Smynthurodes betae
P. texana Geopemphigus sp.
P. vera Baizongia pistaciae
(Pistachio) Forda hirsuta, marginata,
?riccobonii
 Pistacia 327

Geoica utricularia
Paracletus cimiciformis
Slavum lentiscoides, mordvilkoi
Smynthrodes betae

Key to alatae from galls on Pistacia

1 ANT III and IV each usually with only a single apical secondary rhinarium
resembling the primary rhinaria on V and VI (sometimes absent from IV,
or rarely one additional small rhinarium is present on III) . . . . . . . . 2
- ANT III always with at least two secondary rhinaria, and usually with 3
or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2 Primary rhinarium on ANT VI elongate, 2 or more x longer than broad
(Fig. 81 A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aploneura lentisci
- Primary rhinarium on ANT VI oval, like that on V (e.g. Fig. 81B) . .
............................................................. 3
3 R IV+V 0.05-0.08 mm long, less than 0.5 x HT II . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Slavum lentiscoides
- R IV+V 0.08-0.13 mm long, more than 0.5 x HT II . . . . . . . . . . . . . . 4
4 Wax pore plates pigmented and of granular appearance. Forewing veins
Cu 1a and Cu 1b separated at their bases. HT I I 0.20-0.25 mm long . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Slavum esfandiarii
- Wax pore plates unpigmented. Forewing veins Cu l a and Cu l b joined at
their bases. HT II less than 0.18mm long . . . . . . . . . . . . . . . . . . . . . . . 5
5 Head without any distinct median suture. HT II with 4-8 hairs (as well
as the 6 subapical ones). Embryos without mouthparts . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Slavum wertheimae
- Head normal, with a distinct median suture. HT II with 14-17 hairs ( + 6
subapical). Embryos with mouthparts . . . . . . . . . . . Slavum mordvilkoi
6 All antennal segments very thick, with hundreds of small (5-6 µm
diameter) circular secondary rhinaria (Fig. 81C). R IV+V long and
narrow, about 4 x its basal width, with 20 or more accessory hairs
arranged in 4 rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rectinasus buxtoni
- Secondary rhinaria much fewer and usually larger. R IV+V less than
3 x its basal width and with fewer hairs . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ANT III with 2-5 small to very small, round or oval secondary rhinaria
(Fig. 8ID). Marginal wax glands on ABD TERG 1-5 often reduced or
indistinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiphonella cynodonti
- If ANT III has fewer than 6 secondary rhinaria then these are large, occu-
pying at least half of width of segment. ABD TERG 1-5 with or without
marginal wax glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
328 Pistacia

8 ABD TERG 1-5 usually with marginal wax glands . . . . . . . . . . . . . . . 9

- ABD TERG 1-5 without marginal wax glands . . . . . . . . . . . . . . . . . . 18
9 Secondary rhinaria conspicuously ciliated . . . . . . . . . . . . . . . . . . . . . . . 10
- Secondary rhinaria not ciliated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10 ANT PT 3-5 times longer than its basal width; ANT PT/BASE 0.40-0.60
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- ANT PT 1-2 times longer than its basal width; ANT PT/BASE 0.25-0.35
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 ANT III a little longer than ANT VI (incl. PT), with 5-12 secondary
rhinaria arranged in a row. Abdominal wax glands with closely adpressed
facets forming cohesive groups . . . . . . . . . . . . . . Asiphonella dactylonii
- ANT III a little shorter than ANT VI (incl. PT), with 5-14 secondary
rhinaria distributed irregularly, never in a row (Fig. 81E). Abdominal wax
glands with facets loosely grouped, often not touching . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetogeoica foliodentata
12 ANT III more than 2 x ANT VI (incl. PT), and bearing 13-20 narrow
transversely elongate secondary rhinaria. Abdomen with broad dark
transverse bands on ABD TERG 3-6 . . . . . . . . . . . Geopemphigus spp.
- ANT III a little shorter than ANT VI (incl. PT), and bearing 3-5 rounded
or broadly oval secondary rhinaria (Fig. 81F). Dorsal abdomen unpig-
mented or with only faint narrow cross-bands . . . . Aloephagus myersi
13 First tarsal segments usually with two sense pegs between the two long
hairs. ANT III as short as or a little shorter than last antennal segment
(incl. PT); PT about 3 x its basal width (Fig. 81G) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baizongia pistaciae
- First tarsal segments with one sense peg between the two long hairs. ANT
III longer than last antennal segment. ANT PT shorter; less than 2 x its
basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 ABD TERG 2-8 (or 3-8) each with a broad dusky or dark transverse band.
Anal plate not joined to CAUDA. Embryos without mouthparts . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica mimeuri
- At least ABD TERG 2-4 without any distinct dark transverse bands. Anal
plate extending dorsolaterally, the extremities joined with the CAUDA to
form a perianal ring. Embryos with mouthparts . . . . . . . . . . . . . . . . . 15
15 ANT III with only 3-7 large secondary rhinaria, usually in a single row
(e.g. Fig. 81I). ANT IV with 1-2 and V with 0-1 secondary rhinaria (when
antennae 6-segmented). Embryos with spatulate dorsal hairs . . . . . . 16
- ANT III with at least 8 secondary rhinaria, not in a single row, and IV
with at least 3 rhinaria (when antennae 6-segmented; e.g. Fig. 81H).
 Pistacia 329

Embryos with dorsal hairs pointed or (rarely) spatulate (Geoica utricularia

group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 ABD sternite 7 with a narrow, dark, transverse band separate from and
anterior to subgenital plate. ANT III with at least 4 hairs, 8-15 µm long.
Embryos of BL less than 0.7 mm, with a transverse row of 6-8 spatulate
spinopleural hairs between the long marginal hairs on ABD TERG 1-7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica lucifuga group
- ABD sternite 7 without a transverse band. ANT III with 0-2 hairs, 4-9 µm
long. Embryos with BL more than 0.8 mm, and with a transverse row of
at least 10 spatulate spinopleural hairs between the long marginals on each
of ABD TERG 1-7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica setulosa
17 Embryos with 14-30 long very finely pointed spinopleural hairs between
similar marginal hairs on ABD TERG 1-7, and 35-64 similar hairs on anal
plate. Hind tibiae short, usually less than 0.25 x BL. ANT I I I with 6-23
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica harpazi
- Embryos with 6-17 long but acuminate hairs between similar marginal
hairs on ABD TERG 1-7, and 16-50 similar hairs on anal plate. Hind
tibiae often more than 0.25 x BL. ANT III with 7-39 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica utricularia group
18 Antenna 5-segmented. Embryos with elongate hind tarsi . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Fordini 'sp. B' of Koach and Wool (1977)
- Antenna 6-segmented. Embryos with hind tarsi only slightly longer than
fore and mid-tarsi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 ANT VI longer than ANT I I I , with a very large elongate primary
rhinarium occupying almost the whole segment (Fig. 81J) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forda formicaria
- ANT VI not longer than ANT III, with primary rhinarium not especially
large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Secondary rhinaria numerous, small, subcircular. ANT PT reduced to a
very short, almost non-existent stump, much shorter than its basal width
(Fig. 81N) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracletus cimiciformis
- Secondary rhinaria variable but usually transversely oval or quadrate,
with long axis occupying more than half width of segment. ANT PT as
long as or longer than its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 All rhinaria with broad sclerotized rims. ANT III hardly longer than ANT
VI, and bearing 3-6 mainly large secondary rhinaria (Fig. 81O) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smynthurodes betae
- Rhinaria without broad sclerotized rims. ANT III 1.5 or more x ANT VI,
and often with more than 6 secondary rhinaria . . . . . . . . . . . . . . . . . 22
22 HT II with 4-8 dorsal and 2-7 ventral hairs (as well as 6 subapical hairs)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
 Pistacia 331

Fig. 81. Antenna of spring migrant alata (from gall) of A, Aploneura lentisci, B, Slavum lentiscoides,
C, Rectinasus buxtoni, D, Asiphoniella cynodonti, E, Chaetogeoica foliodentata, f, Aloephagus myersi,
G, Baizongia pistaciae, H, Geoica utricularia, I, G. setulosa, J, Forda formicaria, K, F. marginata,
L, F. riccobonii, M, F. hirsuta, N, Paracletus cimiciformis, 0, Smynthurodes betae.
332 Pistacia

- HT II with 1-4 dorsal and 0-3 ventral hairs ( + 6 subapical) . . . . . . 25

23 Antennae short, of total length only about 1.5 x head width across (and
including) eyes or less, with ANT III 1.1-1.2 x HT II. Hairs on ANT III
maximally 15-20 µm, at least as long as diameter of constricted base of
segment. Secondary rhinaria distributed ANT III 6-14, IV 2-4, V 0-3, VI
BASE 0-2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forda sichangensis
- Antennae more than 1.6 x longer than head width across eyes, with ANT
III 1.4-1.9 x HT II. Hairs on ANT III maximally about 10µm or less,
much shorter than diameter of constricted base of segment. Secondary
rhinaria distributed ANT III 15-30, IV 6-12, V 4-12, VI BASE 3-10 (e.g.
Fig. 81K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Longest hairs on ventral side of hind tibia 60-80 µm long. First segment
of middle tarsus with two sense pegs . . . . . . . . . . . . . . . Forda kaussarii
- Longest hairs on ventral side of hind tibia 25-35 µm. First segment of
middle tarsus with only one sense peg . . . . . . . . . . . . . Forda marginata
25 Secondary rhinaria distributed ANT I I I 10-27, IV 5-10, V 2-6 and VI 1-5;
mostly transversely oval, not extending half-way around segment
(Fig. 81L). Embryos with short hairs (about 10 µm long) on ABD TERG
1-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forda riccobonii
- Secondary rhinaria distributed ANT III 7-15, IV 2-6, V 1-4, VI 0-2;
mostly narrow transversely elongate, often extending more than half-way
around segment (Fig. 81M). Embryos with hairs on ABD TERG 1-5 more
than 20 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forda hirsuta

Partial key to alate sexuparae (embryos without mouthparts)

1 Secondary rhinaria with ciliated rims . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Secondary rhinaria with plain rims . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Antennae 6-segmented, with ANT III, IV and V all similar to one another,
each with a single large rhinarium near the apex; sometimes III also bears
another, smaller rhinarium . . . . . . . Aploneura lentisci and Slavum sp.
- Antennae 5- or 6-segmented, ANT III clearly longer than IV and bearing
10-20 rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Antennae 5-segmented, secondary rhinaria distributed III 10-11, IV 0-1.
Abdominal segments 1-6 bearing marginal (lateral) tubercles. ABD TERG
8 with 6-8 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aloephagus myersi
- Antennae 6-segmented, secondary rhinaria distributed III 10-20, IV 1-3.
Abdomen without marginal tubercles. ABD TERG 8 with 2-3 hairs . . . . .
............................................................. 4
4 R IV+V only 0.06-0.07 mm long, less than 0.4 x HT II which is
0.18-0.20 mm. Secondary rhinaria distributed ANT III 10-13, IV 1-3.
Only dorsal wax glands present . . . . . . . . . . . . . . Asiphonella cynodonti
 Pistacia 333

- R IV+V 0.09-0.13 mm long, at least 0.6 x HT II which is 0.13-0.18 mm.

Secondary rhinaria distributed ANT I I I 13-20, IV 1-12. Abdominal
sternites with paired wax glands . . . . . . . . . . . . . . . . . Geopemphigus sp.
5 Primary rhinaria with ciliated rims . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Primary rhinaria with plain rims; only the small accessory rhinaria on VI
sometimes ciliated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6 ANT II bearing 5-7 hairs, and ANT BASE VI with 4-6 hairs. Mesothorax
often with a pair of wax pore plates; abdomen with large lateral and small
medial wax pore plates. R IV+V only 0.6-0.7 x HT II and bearing only
2 (rarely 3 or 4) accessory hairs. Anal plate with a longitudinal double row
of 4 or 5 pairs of long thick hairs in addition to a group of shorter hairs.
First tarsal segments respectively with 4, 4 and 3 hairs. ANT PT/BASE
0.35-0.45 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baizongia pistaciae
- ANT II bearing 6-50 hairs, and ANT BASE VI (or BASE V if antenna
5-segmented) with 9-40 hairs. Mesothorax without wax pore plates;
abdomen without lateral wax pore plates, sometimes with small groups
of pores dorsally on the unpigmented area or posteriorly on the pigmented
parts of the tergites. R IV+V as long as or longer than HT II and bearing
4-10 accessory hairs. Anal plate either bearing only numerous hairs of
similar length or, if w i t h a double row of larger hairs then there are 6 or
7 pairs. First tarsal segments respectively w i t h 3, 3 and 2-3 hairs. ANT
PT/BASE 0.17-0.30 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ABD TERG 8 with numerous (60 + ) scattered hairs similar to those on
anal plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica harpazi
- ABD TERG 8 with 7-20 hairs more or less in a transverse row . . . . 8
8 Anal plate with numerous scattered hairs . . . . . . . . . Geoica utricularia
- Anal plate with 6-7 pairs of long stout hairs in a longitudinal double row
in addition to a group of shorter hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 ABD TERG 5 and 6 bearing scattered hairs like ABD TERG 2-4. Wax
pores present only between ABD TERG 1-3 except in very large speci-
mens. Eongest hair on marginal abdominal sclerites 2-4 is 25-40 µm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica setulosa
- ABD TERG 5 and 6 with only a transverse row of hairs similar to those
on ABD TERG 7 and 8; ABD TERG 2-4 with scattered hairs. Wax pores
commonly present dorsally between ABD TERG 4-7 as well as 1-3.
Longest hair on marginal abdominal sclerites 2-4 is c. 60 µm long . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geoica lucifuga
10 R IV+V 0.7-1.0mm long, about 3 x HT I I . First tarsal segments respec-
tively with 3, 3 and 2 hairs . . . . . . . . . . . . . . . . . . . . Rectinasus buxtoni
- R IV+V 0.13-0.57 mm long, 0.6-1.4 x HT II. First tarsal segments
respectively with 5-12, 5-12 and 4-8 hairs . . . . . . . . . . . . . . . . . . . . . . 11
334 Pithecellobium

11 Antennae 6-segmented. R IV+V 0.13-0.22 mm, shorter than HT II

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- Antennae 5-segmented. R IV+V 0.19-0.60 mm, longer than HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
12 ANT III and IV with numerous rhinaria (52-71 and 21-50 respectively).
Small accessory rhinaria near the primary rhinarium on VI have simple
rims. R IV+V with c. 30 accessory hairs. First tarsal segments with 9 or
more hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- ANT III with 6-25 rhinaria, IV with 1-5. Accessory rhinaria on VI with
ciliated rims. R IV+V with 8-14 accessory hairs. First tarsal segments
respectively with 5, 5 and 4 hairs . . . . . . . . . . . . . . Smynthurodes betae
13 ANT III 1.4-1.9 x ANT IV; R IV+V 0.55-0.70 x HT II, which is
0.30-0.32 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracletus cimiciformis
- ANT III 0.7-0.8 x ANT IV; R IV+V 0.40-0.55 X HT II, which is
0.37-0.47 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracletus donisthorpei
14 HT II very long, 0.70-0.95 mm. R IV+V 0.45-0.60 mm . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Fordini 'sp. B' of Koach and Wool (1977)
- HT II 0.18-0.27 mm, R IV+V 0.19-0.31 mm . . . . . . . . . . . . Forda spp.

PITHECELLOBIUM (= PITHECOLOBIUM) Leguminosae

Pithecellobium dulce Aphis craccivora

P. lucidum Anomalosiphum pithecolobii

PITTOSPORUM Pittosporaceae

Aphis craccivora, gossypii, fabae,

spiraecola
[Greenidea anonae]
Macrosiphum euphorbiae
Myzus antirrhinii, persicae
[Sitobion sp.; Millar, 1991]
Toxoptera aurantii, odinae
(Use key to polyphagous aphids, p. 532)
 Podocarpus 335

PLAGIANTHUS Malvaceae

Plagianthus betulinus Aphis sp.

PLANERA Ulmaceae

Planera aquatica Tetraneura ulmi

PLATANUS Planes Platanaceae

Aphis craccivora, gossypii

Longistigma caryae
Myzus persicae
(Use key to polyphagous aphids, p. 532)

PLATYCARYA Juglandaceae

Kurisakia sinoplatycaryae

PODOCARPUS Podocarpaceae

Host Plant List

Podocarpus alpina Neophyllaphis gingerensis
P. chinensis Neophyllaphis sp. nr
brimblecombei, podocarpi
P. elatus Neophyllaphis brimblecombei,
fransseni
P. engelmannii Neophyllaphis grobleri
P. falcata Neophyllaphis grobleri, viridis
336 Podocarpus

P. gracilior Neophyllaphis grobleri

P. hallii Neophyllaphis totarae
P. henckeli Neophyllaphis grobleri, podocarpi
P. latifolia Neophyllaphis grobleri
P. macrophylla Neophyllaphis podocarpi
P. milanjiana Neophyllaphis grobleri
P. nageia Neophyllaphis podocarpi
P. neriifolia Neophyllaphis podocarpi
P. nivalis Neophyllaphis totarae
P. nubigena Neophyllaphis michelbacheri
P. saligna Neophyllaphis podocarpini
P. spinulosa Neophyllaphis lanata
P. totara Neophyllaphis totarae
P. usumbarensis Neophyllaphis grobleri
Podocarpus sp. ('numphii') Neophyllaphis fransseni

Key to Neophyllaphis spp. on Podocarpus

(This key cannot be applied to oviparae, which are usually winged, except
where stated).
1 Knob of CAUDA globose, about as wide as long . . . . . . . . . . . . . . . . 2
Knob of CAUDA elliptical or elongate, much longer than its maximum
width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Knob of CAUDA with 10-16 hairs, including a single median dorsal hair.
SIPH cone with 2 small hairs. Aptera with eye of 3 facets only . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . grobleri
- Knob of CAUDA with 18-30 hairs, with or without a dorsal hair. SIPH
cone with a ring of 6-8 hairs. Aptera with compound eyes . . . . . . . . 3
3 HT II 0.22-0.32 mm, more than 2.5 x R IV+V. Length of ANT III of
aptera about equal to head width across (and including) eyes . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . podocarpini
- HT II 0.16-0.19 mm, less than 2.5 x R IV+V. Length of ANT III of
aptera about half of head width across eyes . . . . . . . . . . michelbacheri
4 Knob of CAUDA with 16-25 hairs . . . . . . . . . . . . . . . . . . . . . . . . . viridis
Knob of CAUDA with 9 or fewer hairs . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 ANT III of aptera 2.8 or more x ANT IV, and ANT I I I of alata
4.5-5.5 x ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . fransseni
 Polyscias 337

- ANT III of aptera less than 2.6 x ANT IV, and ANT III of alata less
than 3.5 x ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 ANT III of aptera less than 0.26 mm long, less than 0.8 of length of fore
tibia. ANT III of alata less than 0.4mm long, bearing 10-25 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
ANT III of aptera more than 0.26mm long, more than 0.8 of length
of fore tibia. ANT III of alata more than 0.4mm long, bearing 30-58
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 Aptera with many dorsal abdominal hairs arising from pigmented
scleroites of varying size and shape. Pigmented areas of head without a
reticulate pattern and localized, the dorsal and lateral areas being well
separated, with a clear area posteriolaterad to antennal base. Alata with
10-22 seconday rhinaria on ANT I I I , usually not extending to apex of
segment, and ANT IV without rhinaria . . . . . . . . . . . . . . . . . gingerensis
Aptera with dorsal abdominal hairs not based on pigmented scleroites.
Pigmented areas of head with a reticulate pattern, the dorsal and lateral
areas being contiguous, and the lateral pigmentation extending forward
from triommatidium to antennal base. Alata with 20-25 secondary
rhinaria on ANT I I I , distributed over whole length of segment, and
ANT IV often with a few rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . totarae
8 Aptera with ANT III 0.35-0.62 mm long; ANT I I I and ABD TERG
3-5 with longest hairs 9-18 µm long. Alata w i t h ANT III a little longer
than IV+V+VI together, bearing 39-58 secondary rhinaria, and knob
of CAUDA pale and without spinules on proximal half of dorsal
surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brimblecombei/lanata
- Aptera with ANT I I I 0.25-0.38 mm long; ANT III and ABD TERG
3-5 with longest hairs 20-30 µm long. Alata with ANT III usually a
little shorter than IV+V+VI together, bearing 30-44 rhinaria, and knob
of CAUDA entirely pigmented and spinulose . . . . . . . . . . . . podocarpi

POLYSCIAS Araliaceae

Polyscias spp. Aphis spiraecola

Myzus persicae
Toxoptera aurantii
(Use key to polyphagous aphids on p. 532)
338 Populus

POPULUS Poplars Salicaceae

There are about 120 poplar-feeding aphid species, in 24 genera. Although

some species appear to be host-specific, many others are not, and there are no
clear patterns of host specificity related to the subgeneric classification of
Populus. Therefore the poplar-feeding aphids are treated as a whole, and two
rather long keys are provided, one to the free-living aphids and the other to
those that form galls or pseudogalls (e.g. folded leaves or 'leaf nests'). The
gall-formers all belong to subfamily Pemphiginae; of these, only the alatae
from the galls are keyed.

Host Plant List

Populus acuminata Aphis maculatae
(Lanceleaf Cottonwood) Chaitophorus populicola
Mordwilkoja vagabunda
Pemphigus populicaulis,
populitransversus
P. adenopoda Chaitophorus populeti
(Chinese Aspen)
P. alba Chaitophorus longisetosus,
(White Poplar) manaliensis, melanosiphon,
populeti, populeti ssp.
sensoriatus, populialbae,
populialbae ssp. yomefuri
Gootiella alba
Pachypappa ?pseudobyrsa,
vesicalis, warshavensis
[Pemphigus bursarius, protospirae,
vesicarius]
Phloeomyzus passerinii
Phylloxerina populi
Pterocomma [anyangense],
populeum, populeum ssp.
dubium, sinipopulifoliae,
smithiae, yezoense
P. angulata Chaitophorus populicola
Pachypappa pseudobyrsa
P. angustifolia Aphis maculatae
(Narrow-Leaf Cottonwood) Chaitophorus populicola,
populifolii, stevensis
Cornaphis populi
 Populus 339

Pachypappa sacculi
Pemphigus betae, know1toni,
monophagus, populiglobuli,
populitransversus, populivenae
Pterocomma populifoliae,
pseudopopuleum, smithiae
Thecabius gravicornis,
populimonilis
P. balsamifera Aphis maculatae
(Tacamahac or Balsam Poplar) Chaitophorus leucomelas,
populialbae, populicola,
populifolii
Clydesmithia canadensis
Epipemphigus niisimae
Pachypappa sp.
Pemphigus betae, borealis,
monophagus, populicaulis,
populiglobuli, populiramulorum,
populitransversus, populivenae
Phylloxerina salicicola
Pterocomma bicolor, populeum,
populifoliae, [ rufipes]
Thecabius affinis, gravicornis,
populiconduplifolius,
populimonilis
P. x berolinensis Pemphigus bursarius, passeki,
phenax
Thecabius affinis
P. bolleana Chaitophorus populeti, populialbae
Neopemphigus turajevi
Pterocomma kozhuchovae
P. x canadensis - P. x euroamericana
P. candicans = P. balsamifera
P. x canescens Chaitophorus populeti, populialbae
(Grey Poplar) Pachypappa tremulae, vesicalis,
warshavensis
P. cathayana Epipemphigus sanpupopuli
Pemphigus tibetensis
Pterocomma neimongolense,
sanpunum
P. ciliata Chaitophorus kapuri
(Bangikat Poplar) Doraphis populi
Epipemphigus imaicus, marginalis
 340 PopuIus

Pemphigus [ignotus], immunis,

matsumurai, mordvilkoi,
nainitalensis, siphunculata,
[venosus]
Phloeomyzus passerinii
Pterocomma ?populifoliae
P. davidiana Chaitophorus ?dorocolus
Doraphis populi
P. deltoides Chaitophorus leucomelas,
(Eastern Cottonwood) neglectus, populicola,
populicola ssp. patchae,
populifolii, stevensis
Mordvilkoja vagabunda
Pachypappa pseudobyrsa
Pemphigus bursarius,
junctisensoriatus, longicornus,
nortonii, obesinymphae,
populicaulis, populitransversus,
populivenae
Pterocomma smithiae
Thecabius populiconduplifolius
P. deltoides var. occidentalis Chailophorus populicola,
(=P. monilifera) ( = P. sargentii) populicola spp. patchae,
(Plains Cottonwood) populifolii, stevensis
Mordvilkoja vagabunda
Pachypappa pseudobyrsa
Pemphigus betae, bursarius,
junctisensoriatus, nortonii,
. populicaulis, populiramulorum,
populitransversus
Phylloxerina popularia, prolifera
Pterocomma pseudopopuleum,
smithiae, yezoense
Thecabius populiconduplifolius
P. densa [Pachypappa shaposhnikovi
GabridJ
Pemphigus iskanderkuli
[Thecabius luppovae Narzikulov]
P. diversifolia Chaitophorus diversifolia
Lambersaphis pruinosae
P. euphratica Chaitophorus euphraticus,
populicola
Pachypappa sp.
Pemphigus immunis, napaeus
 Populus 341

P. x euroamericana Chaitophorus euphraticus,

leucomelas
Pemphigus populinigrae
Phloeomyzus passerinii
Pterocomma populeum
P. fremontii (incl. var. wislizeni) Chaitophorus populifolii
(Fremont Poplar) Pachypappa ?pseudobyrsa
Pemphigus ?canadensis,
populicaulis, populiramulorum,
populitransversus, populivenae
Phylloxerina popularia
Thecabius populimonilis
P. x generosa Pemphigus sp.
Pterocomma populeum
P. grandidentata Chaitophorus neglectus, stevensis
(Bigtooth Aspen) Pterocomma populifolii, smithiae
P. heterophylla No aphids recorded
(Swamp Cottonwood)
P. koreana Chaitophorus dorocolus, inouyei,
variegatus
Epipemphigus niisimae
Pemphigus borealis, dorocola,
matsumurai
Pterocomma yezoense
Thecabius affinis
P. laurifolia Chaitophorus populeti, populialbae,
tremulae, tremulae ssp. sorini
Pachypappa marsupialis ssp.
lambersi, [shaposhnikovi
Gabrid, 1989]
Pemphigus borealis, laurifoliae,
mongolicus, montanus, sp. nr
iskanderkuli
Phloeomyzus passerinii
Pterocomma populeum ssp. dubium
Thecabius affinis
P. maximowiczii Chaitophorus dorocola, inouye,
(Doro-no-ki) populeti, tremulae ssp.
shantungensis
Doraphis populi
Epipemphigus niisimae
Pachypappa marsupialis ssp.
lambersi
[Pachypappella
342 Populus

orientalis Mordvilko ex
Shaposhnikov, 1955]
Pemphigus borealis, dorocola,
matsumurai, microsetosus,
semenovi
Phloeomyzus passerinii
Pterocomma yezoense
Thecabius affinis, latisensorius,
orientalis
Tuberolachnus salignus
P. monilifera = P. deltoides var. occidentalis
P. nigra (incl. var. italica) Aphis maculatae
(Black Poplar, Lombardy Poplar) Aulacorthum solani
Chaitophorus euphraticus,
leucomelas, nassonowi,
neglectus, populeti, populialbae,
populicola, pruinosae
Pachypappa marsupialis
Pemphigus bursarius, gairi,
immunis, passeki, populi,
populinigrae, protospirae,
spyrothecae, vesicarius
Phloeomyzus passerinii
Phylloxerina populi
Pterocomma bicolor, populeum,
tremulae, yezoense
Stomaphis longirostris
Thecabius affinis
Thecabius (Parathecabius)
lysimachiae
Tuberolachnus salignus
P. occidentalis = P. deltoides var. occidentalis
P. pruinosa Chaitophorus pruinosae
Lambersaphis pruinosae
Pachypappa warshavensis
P. rasumowskyana Pemphigus bursarius, ?populinigrae
P. x robusta Chaitophorus leucomelas,
nassonowi
P. sargentii = P. deltoides var. occidentalis
P. sieboldii Chaitophorus dorocolus, populeti,
populialbae ssp, yomefuri,
tremulae
Doraphis populi
Pachypappa tremulae
 Populus 343

P. simonii Chaitophorus clarus, leucomelas,

tremulae ssp. shantungensis
Pemphigus sinobursarius
Pterocomma neimongolense,
sanpunum, sinipopulifoliae
P. sosnovskii Pemphigus protospirae
P. suaveolens Chaitophorus populeti
Epipemphigus niisimae
Gootiella tremulae
[Pachypappella orientalis
Mordvilko ex Shaposhnikov,
1955]
Pemphigus borealis, dorocolus,
inouye, matsumurai, semenovi,
vesicarius
Phloeomyzus passerinii
Pterocomma populeum, yezoense
Thecabius affinis
P. tadzhikistanica Pemphigus bursarius, montanus,
populi, populinigrae,
protospirae, vesicarius
Phloeomyzus passerinii
P. tianschanica Chaitophorus leucomelas
Pemphigus protospirae
P. tremula (incl. var. davidiana, Chaitophorus populeti, populialbae,
var. villosa) tremulae
(European Aspen) Doraphis populi, populi ssp.
tremulae
Gootiella tremulae
Pachypappa populi, tremulae
Pachypappella lactea
Pemphigus circellatus
Pterocomma tremulae
Stomaphis longirostris
P. tremuloides Aphis maculatae
(Quaking Aspen) Chaitophorus neglectus, nudus,
populicola, populicola ssp.
patchae, stevensis
Fullawaya ontarioensis
Pachypappa pseudobyrsa, rosettei,
sacculi
Pemphigus populicaulis,
populitransversus
Pseudopterocomma canadensis,
hughi
344 Populus

Pterocomma beulahensis, bicolor,

populifoliae, smithiae
P. trichocarpa Chaitophorus populifolii
(Black Cottonwood, Western Clydesmithia canadensis
Balsam Poplar) Pachypappa sp.
Pemphigus betae, monophagus,
populicaulis, populiglobuli,
populivenae
Phylloxerina popularia
Pterocomma smithiae
Thecabius populiconduplifolius,
populimonilis
P. veraecki Pemphigus sp.
P. wislizeni = P. fremontii var. wislizeni
Populus sp. (unidentified) Ceratoglyphina populisucta Zhang
and Zhong, 1985a (= Doraphis
populi?)
Chaitophorus hillerislambersi,
indicus, miyazakii, nodulosus
Longistigma caryae
Neopterocomma populivorum
[Pachypappa pilosa]
Pemphigus [chomoensis],
[cylindricus], [mangkamensis],
napaeus, popularius, similis,
yangcola, [yunnanensis]
Phloeomyzus passerinii ssp.
zhangwuensis Zhang, 1982 (in
Zhang and Zhong, 1982b)
[Prociphilus populi Tao, 1970]
Pterocomma bailangense, medium,
lhasapopuleum

KEY l - Aphids free-living on stems or leaves of Populus

Applies only to adult apterous viviparae unless otherwise stated.
1 Body aleyrodiform (dark, flattened, almost circular, with a complete,
crenulate margin of wax glands). Head of immature alata with a pair of
prominent frontal horns . . . . . . . . . . . . . . . . . . . . . . . . . . Doraphis populi
- Body not aleyrodiform and head of immature alata without a pair of
frontal horns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Body small, pear-shaped (broadest anteriorly), with faceted wax pore
plates on all tergites. Antennae 3-segmented . . . . . . . Phylloxerina spp.
- Body not broadest anteriorly, without wax glands or with wax glands on
ABD TERG 7. Antennae 5-or 6-segmented . . . . . . . . . . . . . . . . . . . . . . 3
 Populus 345

3 Eyes with only 3 facets. ABD TERG 7 with a pair of large wax pore plates.
Antennae with spinules in a reticulate pattern . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phloeomyzus passerinii
- Eyes multifaceted. Abdomen without wax glands. Antennae not orna-
mented with a reticulate pattern of spinules . . . . . . . . . . . . . . . . . . . . . . 4
4 CAUDA tongue- or finger-like, tapering, much longer than its basal width
............................................................. 5
- CAUDA knobbed, rounded or triangular . . . . . . . . . . . . . . . . . . . . . . . . 7
5 Antennal tubercles undeveloped. SIPH and CAUDA black. ANT III with
15-22 secondary rhinaria over distal 0.8 of length. Tibiae with numerous
long, fine hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis maculatae
- Antennal tubercles well developed. SIPH and CAUDA mainly pale. ANT
III with 1-9 secondary rhinaria on basal 0.4. Tibiae with short hairs
............................................................. 6
6 Antennal tubercles w i t h inner faces approximately parallel (Fig. 42J).
ANT III with a single rhinarium near base. SIPH without an apical zone
of polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
- Antennal tubercles with inner faces clearly divergent (Fig. 42K). ANT III
with 2-9 rhinaria. SIPH with a subapical zone of polygonal reticulation
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
7 Very large aphid with ANT PT/BASE 0.6 or less. SIPH pores placed on
broad dark hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Small or large aphid with ANT PT/BASE more than 0.6. SIPH truncate
conical (with reticulate sculpturing) or tubular, or merely pores, or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
8 Rostrum much longer than body, which is elongate oval. ANT PT/BASE
about 0.1, with PT broad and rounded . . . . . . . Stomaphis longirostris
- Rostrum much shorter than body, which is broadly oval. ANT PT/BASE
0.3-0.5, with PT tapering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Dorsal abdomen with a large dark spinal tubercle. BL usually less than
5.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberolachnus salignus
- Dorsal abdomen without a tubercle. BL usually more than 5.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 HT II less than 2.2 x HT I. ANT III with longest hairs much longer than
diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- HT II more than 2.2 x HT I. ANT III with longest hairs shorter than
diameter of segment . . . . . . . . . . . . . . . . . . . . . Longistigma xizangensis*
11 SIPH in form of small truncate cones or short cylinders (usually at least
0.5 x basal width), with reticulate sculpturing at least distally. CAUDA
knobbed or rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
346 Populus

Fig. 82. A, hind tibia of apterous vivipara of Chaitophorus populeti with scent plaques; CAUDA of aptera
of B, Ch. nassonowi, C, Ch. populicola and D, Ch. populialbae; HT I of E, Ch. populialbae and F, Ch. tremulae.

- SIPH without reticulate sculpturing; either in form of flat cones (less

than 0.5 x basal width), or tubular, or pore-like, or absent. CAUDA
rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
12 Hind tibia with 1-22 scattered small scent plaques ('pseudorhinaria';
Fig. 82A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Hind tibia without any scent plaques (except in oviparae) . . . . . . . . 15
13 ANT PT/BASE 0.9-1.4. CAUDA variable but often nearly triangular,
with a rounded apex and almost without any constriction (Fig. 82B)
......................................... Chaitophorus nassonowi
- ANT PT/BASE 1.9-3.1. CAUDA always with a distinctly knobbed apex
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Antennal hairs with finely pointed apices. (Alata with secondary rhinaria
not in a row, distributed III 10-32, IV 0-8, V 0-2) ................
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus populeti
- Antennal hairs rather stiff, mostly with blunt or bifurcate apices. (Alata
with secondary rhinaria more-or-less in a row, distributed III 10-20,
IV 0-4, V 0-1) . . . . . . . . . . . . . . Chaitophorus populeti ssp. sensoriatus
15 CAUDA rounded, without any trace of a constriction (e.g. Fig. 82C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- CAUDA with a constriction delimiting an apical knob (Fig. 82D) ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
 Populus 347

16 ABD TERG 1-7 usually all free, with separate spinopleural and marginal
sclerites. Tibiae with pale middle sections . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus miyazakii *
- ABD TERG 2-7 wholly sclerotized, fused into a solid tergum. Tibiae
wholly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 ABD TERG 2-7 each with only 3-5 spinal hairs (plus marginals), the
pleural regions being devoid of hairs . . . . . . . . . . . Chaitophorus nudus
- ABD TERG 2-7 each with pleural as well as spinal and marginal hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Abdominal tergum evenly spinulose or nodulose . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus nodulosus
- Dorsal cuticle without evident spinules or nodules . . . . . . . . . . . . . . . 19
19 Many of dorsal abdominal hairs blunt or w i t h square or furcate apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus populicola
- All dorsal abdominal hairs long, fine and acute . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus populicola ssp. patchae
20 First tarsal segments usually with 5 hairs (e.g. Fig. 82E; rarely 6 or more
on some tarsi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- First tarsal segments usually with 7 hairs (e.g. Fig. 82F; rarely 6 on some
tarsi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
21 ABD TERG 1-6 fused, wholly dark, distinctly ornamented with minute
spinules, and with numerous smaller pointed hairs between the very long
spinal, pleural and marginal hairs on each segment, which have usually
pointed but sometimes blunt or bifurcate apices (Fig. 83A) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus longisetosus
- Dorsal abdomen pale, or with separate dark segmental bands; more-or-
less smooth or ornamented with blunt nodules or spinules, or polygonal
reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 ANT PT/BASE 1.5 or less (fundatrix has ANT PT/BASE 0.6-0.9) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus pruinosae*
- ANT PT/BASE 1.7 or more (1.3 or more in fundatrix) . . . . . . . . . . 23
23 R 1V+V shorter than HT II, with 2 accessory hairs . . . . . . . . . . . . . . 24
- R IV+V as long as or a little longer than HT II, with 2-4 accessory
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
24 Spinopleural hairs on ABD TERG 2-7 very short, maximally about
20 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus clarus
- Spinopleural hairs long, maximally 100-150 µm . . . . . . . . . . . . . . . . . 25
25 ABD TERG 2-6 with weakly spinulose ornamentation, and long thick hairs
348 Populus

Fig. 83. Hairs in spinal region of ABD TERG 3 of aptera of A, Chaitophorus longisetosus, B, Ch. inouyei,
C, Ch. variegatus, D, Ch. hillerislambersi, E, Ch. tremulae, F, Ch. euphraticus, 6, Ch. stevensis, H, Ch. neglectus.

maximally 150 µm. ANT III with 3-9 hairs, the longest 1.7-2.2 x basal
diameter of segment . . . . . . . . Chaitophorus populialbae ssp. yomefuri
- ABD TERG 2-6 with weakly to strongly nodulose ornamentation, and
hairs up to 120 µm long (Fig. 83F). ANT III with 6-13 hairs, the longest
0.8-1.7 x basal diameter of segment . . . . . . Chaitophorus euphraticus
26 ANT PT/BASE 3.3-4.2. SIPH dark .... Chaitophorus melanosiphon
- ANT PT/BASE 1.7-3.2. SIPH pale . . . . . . . Chaitophorus populialbae
27 Empodial hairs enlarged and flattened, with expanded apices. Abdominal
tergum pale, with long, pointed or spine-like spinal, pleural and marginal
hairs interspersed with smaller hairs, some of which are much smaller
(0.25 x length of longest) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Empodial hairs setaceous, if somewhat flattened then the abdominal
tergum is dark and/or the spinopleural hairs are often blunt or furcate
and/or their variation in size is much less . . . . . . . . . . . . . . . . . . . . . . 30
28 ABD TERG 2-6 with smallest (accessory) hairs mainly thick and spine-
like, and very numerous (30-40 per segment; Fig. 83B) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus inouye
 Populus 349

- ABD TERG 2-6 with accessory hairs finer and less numerous (less than
25 per segment; e.g. Fig. 83C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 Longest hairs on ANT III 3.5-4.9 x basal diameter of segment. ANT PT/
BASE 3.5-4.5. ABD TERG 1-7 mostly smooth, without ornamentation
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus variegatus
- Longest hairs on ANT III 1.7-2.5 x basal diameter of segment. ANT
PT/BASE 2.7-3.4. ABD TERG 1-7 with weak squamous or nodulose
ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus indicus*
30 ABD TERG 1 frequently fused with ABD TERG 2-6 (examine several
specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
- ABD TERG 1 almost always free from ABD TERG 2-6 . . . . . . . . . 37
31 R IV+V 0.17-0.20mm, 1.2-1.4 x HT I I . Abdominal tergum smooth,
bearing mostly long hairs w i t h bifurcate apices (Fig. 83D) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus hillerislambersi
- R IV+V 0.07-0.14 mm, 0.7-1.2 x HT II. Abdominal tergum usually
ornamented with spicules or nodules . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 Dorsal ornamentation consisting of close, irregular rows of posteriorly-
directed, bluntish spinules, often united basally to form partial transverse
rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3
- Dorsal ornamentation consisting of rather evenly distributed separate
roundish nodules (e.g. Fig. 83E; sometimes weakly developed) . . . . 35
33 Dorsum pale. ANT PT/BASE 1.5-2.4 . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus tremulae ssp. sorini
- Dorsum usually deeply pigmented, sometimes dark only on head and ABD
TERG 8. ANT PT/BASE 2.2-3.2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34 Longest hairs on ANT III 1.7-3.2 x basal diameter of segment . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus tremulae s. str.
- Longest hairs on ANT III with wavy, attenuate apices, 3.0-4.5 x basal
diameter of segment . . . . . . Chaitophorus tremulae ssp. shantungensis
35 Hairs on each of ABD TERG 1-7 very variable in size and form, arranged
more or less in a single row (Fig. 83G; sometimes with 1-2 smaller hairs
out of line). ABD TERG 8 with 7-12 hairs. (Alata with SIPH dark)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus stevensis
- Hairs on each of ABD TERG 1-7 arranged in two irregular rows, the
anterior row consisting of smaller hairs which are sometimes absent in
spinal region (e.g. Fig. 83H). ABD TERG 8 usually with more than 12
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 Head and ANT I-II often dark, and tergum often with a pair of dark
longitudinal pleural stripes. Most of long spinal and pleural hairs on ABD
350 Populus

TERG 1-7 with pointed or blunt apices, with a minority expanded or

bifurcate apically (Fig. 83H). (Alata with SIPH dark) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus neglectus
- Head, ANT I-II and tergum usually pale. If with dark pigmentation then
there are rarely complete pleural stripes, and the long spinal and pleural
hairs on ABD TERG 1-7 mostly have expanded or bifurcate apices. (Alata
with SIPH pale) . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus populifolii
37 ABD TERG 1-6 with largest spinal and pleural hairs apically bifurcate
or blunt, marginal hairs pointed. R IV+V 0.7-0.8 x HT II (tentative
characters for aptera, which has not been seen by authors) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus dorocolus
- ABD TERG 1-6 with largest spinal and pleural hairs usually pointed.
R IV+V 0.8-1.2 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 Dorsal cuticle mainly dark, leaving a narrow pale spinal stripe . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus kapuri
- Dorsal cuticle either pale or with dark pleural longitudinal stripes leaving
broad pale spinal and marginal areas . . . . . . Chaitophorus leucomelas
39 SIPH as pores or small flat cones, or absent . . . . . . . . . . . . . . . . . . . 40
- SIPH t u b u l a r , longer than basal width, often swollen . . . . . . . . . . . . 44
40 ANT PT/BASE less than 1; PT w i t h o u t hairs except those at apex. SIPH
as small cones or craters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
- ANT PT/BASE more than 1; PT bearing hairs along length. SIPH pore-
like or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
41 Body sparsely hairy, without spinal or marginal tubercles. R IV with
4-6 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . Lambersaphis pruinosae
- Body densely hairy, with numerous spinal and marginal tubercles. R IV
with 22-26 accessory hairs . . . . . . . . . . . Neopterocomma populivorum
42 Body densely covered in fine hairs. ANT PT/BASE less than 1.5. Well-
developed lateral tubercles on ABD TERG 2-7. SIPH absent . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fullawaya ontarioensis
- Body hairs sparse, especially dorsally. ANT PT/BASE more than 1.5.
Lateral abdominal tubercles absent. SIPH as pores . . . . . . . . . . . . . . 43
43 ABD TERG 1-6 all fused together and fully pigmented. Body hairs not
spine-like . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudopterocomma canadensis
- ABD TERG 1-6 not or hardly pigmented, or with only narrow transverse
intersegmental bars. Body hairs spine-like . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudopterocomma hughi
44 SIPH of most specimens vasiform (swollen on basal half; Fig. 84A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma smithiae
 Populus 351

- SIPH either almost cylindrical or swollen on distal half . . . . . . . . . . 45

45 ANT II with 8-12 hairs. ANT III (aptera) with 4-79 secondary rhinaria,
and ANT IV with 0-9 . . . . . . . . . . . . . . . . . . . . . . Pterocomma tremulae
- ANT II with 4-6 hairs. ANT III with 0-26 secondary rhinaria, ANT IV
never with any . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
46 ANT VI BASE with 6-10 long fine hairs (plus 2-3 short ones) . . . 47
- ANT VI BASE with 1-3 long fine hairs (plus 2-6 short ones) . . . . 49
47 ANT III (aptera) with 14-26 secondary rhinaria . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma kozhuchovae
- ANT III always without secondary rhinaria . . . . . . . . . . . . . . . . . . . . . 48
48 ABD TERG 1-7 each with a pair of large dark quadrate patches
(sometimes reduced or absent on 1-3 or 1-4). Longest hairs on ANT III
100-160 µm, on hind tibia 100-200 µm . . . . . . Pterocomma populeum
- ABD TERG 1-7 without markings or w i t h only lightly pigmented patches
on 6-7 plus scattered small sclerites on 1-5. Longest hairs on ANT III
50-70 µm, on hind tibia 60-80µm . . . . Pterocomma pseadopopuleum
49 ANT PT/BASE of most specimens less t h a n 1.4 . . . . . . . . . . . . . . . . 50
- ANT PT/BASE usually more than 1.4 ( N . B . not applicable to f'unda-
trices) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
50 ANT I with 13-17 hairs. CAUDA with 49-55 hairs, anal plate with 85-98
hairs. SIPH cylindrical . . . . . . . . . . . . . . . . . . . . Pterocomma sanpunum
- ANT I with 8-13 hairs. CAUDA with 24-38 hairs, anal plate with 25-64
hairs. SIPH usually swollen on distal half . . . . Pterocomma yezoense
51 CAUDA with more than 50 hairs . . . . . Pterocomma lhasapopuleum*
- CAUDA with 24-36 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
52 ABD TERG 1 with a single pair of large sclerites, or a band interrupted
in midline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma bailangense
- ABD TERG 1 at most with scattered or paired groups of smaller sclerites
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
53 ANT III (aptera) with 0(rarely)-25 secondary rhinaria. ANT PT
0.15-0.28 mm long. (Alata with 35-50 secondary rhinaria on ANT III and
0-7 on IV) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma populifoliae
- ANT III without rhinaria. ANT PT 0.22-0.36 mm long. (Alata with 12-26
secondary rhinaria on ANT III, 0-2 on I V ) . . . . . . . . . . . . . . . . . . . . 54
54 SIPH usually distinctly swollen (Fig. 84B), 0.30-0.50 mm long, 1.1-1.6 X
PT, which is 0.26-0.36 mm long. CAUDA brown, concolorous with anal
plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma bicolor
352 Populus

Fig. 84. Right SIPH of aptera of A, Plerocomma smithiae, B, Pt. bicolor and C, Pt. beulahense.

- SIPH usually almost cylindrical (Fig. 84C), 0.21-0.28 mm long, 0.8-1.0 x

PT, which is 0.22-0.32 mm long. CAUDA almost black, usually clearly
darker than anal plate . . . . . . . . . . . . . . . . . . . . Pterocomma beulahense

KEY 2 - Aphids forming galls or pseudogalls (e.g. 'leaf nes(s') on poplars

The main key below can be applied only to adult alatae produced in galls or
pseudogalls on Populus. Usually alatae are produced in large numbers in galls
in late spring or early summer to fly to secondary host plants. These alatae
are variously termed spring migrants, emigrants, alate fundatrigenia or funda-
trispuriae. A few species (Cornaphis populi and four species of Pemphigus)
are monoecious (without host alternation). In these, the galls mature in late
summer/autumn and contain numerous alate sexuparae, easily recognized in
cleared microscope preparations by the absence of mouthparts from their
embryos. The key includes the sexuparae of these species, as they are produced
within the galls, but not the sexuparae of heteroecious species, as these are
produced on the secondary host.
Separation of the species of Pemphigus is difficult, and that part of the
key (couplets 24-56) is rather tentative. However, many specimens are usually
available from each gall, so that individual variation in the sample can be
assessed, and tentative identifications can be checked against the illustrations
(Fig. 131) and descriptions of galls in the systematic section of the book. The
complete life cycle of many Pemphigus species is still unknown, so it is likely
that there are many still unrecognized species, as well as synonyms among
the names in current use.
Sometimes aphids that are not gall formers may be found in old galls.
Phylloxerina populariae, for example, regularly inhabits old galls of Thecabius
populiconduplifolius.
 Populus 353

In three species of Pemphiginae, Clydesmithia canadensis, Gootiella alba

and Pemphigus siphunculatus, no alatae have been found even in apparently
mature galls. The complete life cycle of these species is not yet known. The
apterae (apterous fundatrigeniae, not fundatrices) from the galls key as
follows.

1 Abdomen without wax glands . . . . . . . . . . . . . Clydesmithia canadensis

Abdomen with large wax glands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 BL 3.3-4.3 mm. Legs pale. HT II short and thick, less than twice as long
as claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gootiella alba
BL 1.9-3.4mm. Legs dark. HT II normal, more than twice as long as
claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus siphunculatus

Key to alatae from galls

1 Forewing with media usually once-branched. If unbranched then there are
3-8 secondary rhinaria on ANT I I I , 1-3 on IV, 0-2 on V and 0 on VI
BASE and R IV+V is more than 0.6 x HT I I . . . . . . . . . . . . . . . . . . . 2
- Forewing with media unbranched. Antenna with more rhinaria, and/or
R IV+V less than 0.6 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
2 Primary rhinaria on ANT V and VI enlarged and of irregular shape
(Fig. 85A); that on VI elongate, more than 50 µm long. Embryos with
enlarged claws on hind legs . . . . . . . . . . . . . . . . . . . . . Gootiella tremulae
- Primary rhinaria on ANT V and VI rounded or oval, that on VI not longer
than wide. Hind legs of embryos with normal claws . . . . . . . . . . . . . . 3
3 SIPH pores very large, with diameter much greater than that of hind tibia
at midpoint, and with a completely pigmented rim (Fig. 85B) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappella lactea
- SIPH pores absent or small and inconspicuous, with diameter less than
that of hind tibia at midpoint, and with pigmented rim partial (e.g.
Fig. 85C) or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Membrane of forewing with hairs (Fig. 85D). Genital plate with more than
10 hairs on anterior half. ABD TERG 8 usually without wax pore plates,
or with only small groups of facets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Membrane of forewing without hairs. Genital plate with 6-12 hairs on
anterior half. ABD TERG 8 usually with a pair of well-developed wax
pore plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5 ABD TERG 1-7 with well-defined marginal wax pore plates. (SIPH pores
present, CAUDA with 15-30 hairs, and secondary rhinaria on ANT III
10-11, on IV 2-4) . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa vesicalis
- ABD TERG 1-7 without marginal wax pore plates or if present they are
ill-defined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
354 Populus

Fig. 85. A, ANT V and VI of emigrant alata of Gootiella tremulae; B, SIPH and surrounding wax gland
of alata of Pachypappella lactea; C, SIPH of alata of Pachypappa populi; D, hairs on forewing membrane
of Pachypappa vesicalis; ANT III of emigrant alata of E, P. warshavensis and F, P. pseudobyrsa;
G, front of head of alata of Cornaphis populi; ANT VI of emigrant alata of H, Mordvilkoja vagabunda,
I, Epipemphigus niisimae, J, E. sanpopuli, K, E. imaicus.

6 CAUDA with more than 40 hairs. SIPH pores present. Secondary rhinaria
on ANT III 8-13, on IV 2-4 . . . . . . . . . . . . . . . . . . Pachypappa populi
- CAUDA with less than 30 hairs. SIPH pores absent. Secondary rhinaria
on ANT III 5-10, on IV 0-3 . . . . . . . . . . . . . . . . . . Pachypappa sacculi
7 R IV+V 0.60-0.75 x HT II. Media sometimes unbranched . . . . . . . . 8
- R IV+V 0.45-0.55 x HT II. Media always once-branched ......... 9
 PopuIus 355

8 ANT PT/BASE 0.08-0.11. ANT I I I with 4-6 secondary rhinaria with

thick sclerotic rims, often confined to distal half of segment (Fig. 85E)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa warshavensis
- ANT PT/BASE 0.17-0.21. ANT I I I with 5-8 secondary rhinaria with
narrow sclerotic rims, extending onto proximal half of segment (Fig. 85F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa pseudobyrsa
9 ABD TERG 8 with 14-25 hairs. CAUDA with only 3-6 hairs . . . . 10
- ABD TERG 8 with 5-11 hairs. CAUDA with 8-25 hairs . . . . . . . . . 11
10 R IV+V without accessory hairs . . . . . . Pachypappa marsupialis s. str.
- R IV+V with 2-4 accessory hairs Pachypappa marsupialis ssp. lambersi
11 R IV+V usually without any accessory hairs. ANT II with 9-14 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa tremulae
- R IV+V with usually 1-4 accessory hairs. ANT I I with 5-11 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa rosettei
12 ANT PT/BASE greater than 1, w i t h PT bearing 2-3 clear spots, each with
a small, dagger-like stipule (Fig. 85H) . . . . . . . Mordvilkoja vagabunda
- ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Head w i t h median ocellus mounted on a distinct f r o n t a l prominence
(Fig. 85G) (Secondary rhinaria small and oval; on ANT I I I 3-5, on IV 0-2
near apex. Embryos without mouthparts) . . . . . . . . . Cornaphis populi
- Head without a distinct frontal prominence. (Sensoriation of antennae not
as above, embryos with or without mouthparts) . . . . . . . . . . . . . . . . . 14
14 Secondary rhinaria with ciliated (or 'dotted') rims . . . . . . . . . . . . . . . 15
- Secondary rhinaria without ciliated rims . . . . . . . . . . . . . . . . . . . . . . . . 18
15 ANT III and IV each with a single subapical secondary rhinarium, like the
primary rhinaria on V and VI . . . . . . . . . . . . . . Neopemphigus turajevi
- ANT III with 4 or more secondary rhinaria, IV with 2 or more and V with
0-1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 ANT PT normal, about as wide as basal articulation of V I , and bearing
5 hairs near apex (Fig. 85I) . . . . . . . . . . . . . . . . . Epipemphigus niisimae
- ANT PT thickened, broader than basal articulation of VI, bearing 8 or
more hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 ANT PT much enlarged, bearing 20-30 hairs (Fig. 85J). Primary
rhinarium on VI oval and of normal size . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epipemphigus sanpupopuli
- ANT PT bearing 7-14 hairs. Primary rhinarium on VI large and of
irregular shape (Fig. 85K) . . . . . . . . . Epipemphigus imaicus/marginalis
356 Populus

Fig. 86. ANT III of emigrant alata of A, Parathecabius latisensorius and B, Thecabius affinis; antenna of
emigrant alata of C, Pemphigus matsumurai and D, P. populi.

18 Secondary rhinaria broad and strap-like, often almost encircling antenna

and with gaps in between rhinaria mostly much narrower than the rhinaria
themselves, so that more than 50% of total surface area of antenna is
sensoriated (e.g. Fig. 86A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
- Secondary rhinaria narrower (e.g. Fig. 86B), or if they are broader than
gaps in between them (e.g. Fig. 86C) then they do not almost completely
encircle antenna (thus always leaving more than 50% of total surface area
of antenna unsensoriated) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
19 ANT BASE VI usually without secondary rhinaria. (ANT III with 15-26,
IV with 8-11, V with 9-11, VI with 0-1) . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parathecabius gravicornis
- ANT BASE VI with 6-14 secondary rhinaria . . . . . . . . . . . . . . . . . . . 20
20 Secondary rhinaria on ANT III 20-41 (Fig. 86A), IV 10-13, V 10-14,
BASE VI 9-14 . . . . . . . . . . . . . . . . . . . . . . . . . Parathecabius latisensorius
- Secondary rhinaria on ANT III 13-21, IV 4-10, V 5-9 and BASE VI 6-12
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parathecabius lysimachiae
 Populus 357

21 R IV with usually 1-6 small accessory hairs (rarely with none) and first
tarsal segments with 2-6 hairs (usually at least 3 on most tarsi). Secondary
rhinaria very narrow, slit-like (Fig. 86B), with 2-9 on ANT BASE VI
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Thecabius affinis/populiconduplifoliae
- R IV usually without accessory hairs, sometimes with 1-4 (but one of the
3 apical pairs is sometimes displaced to a more proximal position). First
tarsal segments usually with 3 hairs on fore tarsus and 2 hairs on mid-
and hind tarsi. If more, then ANT BASE VI is without secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 SIPH pores absent and embryos with mouthparts (the curled stylets are
especially evident through wall of abdomen) . . . . . . . . . . . . . . . . . . . . 23
- Either SIPH present as small pores or embryos without mouthparts
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
23 R IV+V broadly triangular, less than 2 x its basal width. (Secondary
rhinaria on ANT I I I 5-19, IV 3-7, V 3-4) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . Thecabius populimonilis (fundatrigeniae)
- R IV+V acute, more than 2 x its basal width . . . . . . . . . . . . . . . . . . . 24
24 Secondary rhinaria present on ANT VI. (On III 10-18, IV 4-7, V 3-7,
BASE VI 3-7) . . . . . . . . . . . . . . . . . . . . Pemphigus napaeus/iskanderkuli
- Secondary rhinaria absent from ANT VI . . . . . . . . . . . . . . . . . . . . . . . 25
25 Secondary rhinaria strap-like, parallel-sided (Fig. 86C), many of them
extending more than half-way around antenna, with 2-5 on ANT V
(III 8-13, IV 2-5, V 2-5). ABD TERG 8 with one large wax pore plate
extending more than half-way across tergite, and including 4 hairs . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus matsumurai
- Secondary rhinaria transversely elongate oval, not parallel-sided, extend-
ing up to half-way around antenna but not more, with 0-2 on ANT V
(e.g. Fig. 86D). ABD TERG 8 with a pair of small wax pore plates or a
single median wax pore plate including only 2 hairs . . . . . . . . . . . . . 26
26 First tarsal segments almost always with 2 hairs, rarely with 3. HT II
0.17-0.21 mm long. Secondary rhinaria on ANT III 3-9, IV 1-3,
V 0-2 (Fig. 86D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populi
- First tarsal segments usually with 3-4 hairs on at least some tarsi. HT II
0.20-0.25 mm long. Secondary rhinaria on ANT III 6-11, IV 1-4, V 0-2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus vesicarius
27 Embryos without mouthparts. Secondary rhinaria always absent from
ANT V and VI. SIPH pores absent. (Sexuparae of monoecious species,
produced in the gall) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Embryos with mouthparts. Secondary rhinaria usually present on ANT
V and VI. SIPH present as small pores . . . . . . . . . . . . . . . . . . . . . . . . 31
358 Populus

28 First tarsal segments usually all with 3-5 hairs, rarely with only 2. Spinal
wax pore plates often present on all abdominal tergites . . . . . . . . . . 29
- First tarsal segments most commonly with 3 (-4) hairs on fore tarsus and
2 (-3) hairs on mid- and hind tarsi. Spinal wax pore plates usually absent
from ABD TERG 4-7. (Secondary rhinaria distributed III 3-6, IV 1-3)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
29 R IV+V about 0.08 mm long, slightly shorter than ANT V. Secondary
rhinaria on ANT III 4-7, on IV 1-4 . . . . . . . Thecabius populimonilis
- R IV+V 0.09-0.11 mm long, as long as or slightly longer than ANT V.
Secondary rhinaria on ANT III 6-10, on IV 2-5 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus mongolicus
30 R IV+V usually without accessory hairs. CAUDA with 4-6 hairs . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus monophagus
- R IV+V usually with 1-4 accessory hairs, rarely with none. CAUDA with
2-4 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus spyrothecae
31 ANT BASE VI usually without separate secondary rhinaria (rarely with
1, in which case there are usually none on V) . . . . . . . . . . . . . . . . . . . 32
- ANT BASE VI usually with 2 or more secondary rhinaria separate from
the primary rhinarium (rarely with 1 only, in which case there is usually
1 or more on V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
32 ANT VI with an enlarged primary rhinarium (due to coalescence with
secondary rhinaria), with islands, like that on V . . . . . . . . . . . . . . . . 33
- ANT VI with primary rhinarium of normal size, no longer than wide
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
33 ANT VI with primary rhinarium 0.07-0.10 mm long, occupying more
than half of length of BASE VI (Fig. 87E), or with a separate secondary
rhinarium immediately adjacent to it. Secondary rhinaria on III 3-9, IV
1-3, V 0(-1) and BASE VI 0(-1) . . . . . . . . . . . . . Pemphigus knowltoni
- ANT VI with primary rhinarium less than 0.07 mm long, occupying less
than half of length of BASE VI (Fig. 87F). Secondary rhinaria on III 2-7,
IV 0-3, V 0(-1), BASE VI 0 . . . . . . . . . . Pemphigus populitransversus
34 ANT V with primary rhinarium very large, more than 0.04mm long
and more than half encircling segment, often with small round islands
(Fig. 87G). (Secondary rhinaria on III 7-9, IV 2-4, V 0-1, BASE VI 0)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus immunis
- ANT V with primary rhinarium either like that on VI or, if enlarged,
broader than its axial length which is less than 0.03 mm . . . . . . . . . 35
35 First tarsal segments with two spine-like hairs between the 2 long apical
hairs. ABD TERG 8 usually without a wax pore plate. ANT PT more than
2 x its basal width. (Secondary rhinaria on III 10-14, IV 2-3, V 0(-1),
BASE VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus microsetosus
 Populus 359

- First tarsal segments with 0-1 spine-like hairs between the two long apical
hairs. ABD TERG 8 usually with wax pore plates. ANT PT less than
2 x its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 R IV+V about 0.1 mm, more than 0.55 x HT II. Rhinaria on ANT III
9-13, IV 3-6, V 0-1, BASE VI 0 . . . . . . . . . . . . Pemphigus circellatus*
- R IV+V 0.07-0.085 mm, less than 0.55 x HT II. Rhinaria on ANT III
5-11, IV 1-4, V 0-3, BASE VI 0(-1) . . . . . . . . . . . . . Pemphigus betae
37 Total antennal length 0.9mm or more, about 0.5 x BL . . . . . . . . . . 38
- Total antennal length less than 0.9 mm, less than 0.45 x BL . . . . . 39
38 Secondary rhinaria on ANT I I I 19-27, on IV 6-7, V 9-12, VI BASE 10-12
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus longicornus
- Secondary rhinaria on ANT I I I 15, IV 4, V 4, VI 5 (one specimen only)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus nainitalensis*
39 R IV+V 0.10-0.12 mm long, w i t h R IV straight-sided (Fig. 87A), usually
more than 0.55 X HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
- R IV+V 0.07-0.10 mm long, R I V with sides slightly convex (e.g.
Fig. 87B), usually less than 0.55 x HT I I . . . . . . . . . . . . . . . . . . . . . . . 41
40 Secondary rhinaria narrow, s l i t - l i k e , averaging 8 µm or less in width,
extending almost to base of ANT I I I , with most proximal rhinarium
usually basad to the small tooth on inner side of III near base (Fig. 87C);
on I I I 12-18, IV 3-7, V 4-7, VI BASE 4-7 . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populinigrae
- Secondary rhinaria broader, averaging more than 10µm in width, with
most proximal rhinarium on ANT III distal to tooth (Fig. 87D); on
III 10-14, IV 2-5, V 2-4, VI BASE 2-8 . . . . . Pemphigus protospirae
41 Primary rhinarium on ANT V usually much enlarged, its axial length
35 µm or more, with small islands (Fig. 87H). Secondary rhinaria on
ANT I I I 6-12, IV 2-3, V 0-3, VI BASE 2-5 . . . . . Pemphigus phenax
- If primary rhinarium of ANT V is enlarged then its axial length is less than
35 µm and the antennal sensoriation is different . . . . . . . . . . . . . . . . . 42
42 Total number of secondary rhinaria on ANT V+VI BASE (both antennae)
24 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
- Total number of secondary rhinaria on ANT V+VI BASE (both antennae)
23 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
43 ANT VI BASE slightly shorter than HT I I . (Secondary rhinaria on ANT
III 11-19, IV 4-8, V 4-10, VI BASE 6-11) . . . . . Pemphigus trehernei
- ANT VI BASE slightly longer than HT II . . . . . . . . . . . . . . . . . . . . . . 44
44 ABD TERG 8 with 2-3 hairs. (Secondary rhinaria on I I I 13-19, IV 4-5,
V 6-9, VI BASE 6-10) . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus borealis
- ABD TERG 8 usually with 3-5 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 45
360 Populus

Fig. 87. R IV+V of alata of A, Pemphigus populinigrae and B, P. bursarius; ANT III of emigrant alata of
C, P. populinigrae and D, P. protospirae, with small tooth on inner side arrowed; ANT VI of
E, P. knowltoni and F, P. populitransversus; ANT V and VI of G, P. immunis, H, P. phenax and
I, P. passeki; basal part of ANT III of J, P. populicaulis and K, P. laurifoliae.

45 Secondary rhinaria on III 9-16, on IV 3-6, V 3-8, VI BASE 5-11

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus nortonii
- Secondary rhinaria on III 16-20, on IV 4-5, V 5-6, VI BASE 5-9 ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus yangcola
46 ANT III shorter than ANT IV+V together in most specimens . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
- ANT III usually about the same length as, or longer than, ANT IV+V
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
47 ANT PT finger-like, 2 or more x its basal width. R IV+V 0.09-0.10 mm
long. Secondary rhinaria on ANT III 8-12, on IV 3-4, V 1-4, VI BASE
3-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus plicatus
- ANT PT stubby, less than 1.5 x its basal width. R IV+V 0.07-0.09 mm
long. Secondary rhinaria on ANT III 7-12, IV 3-6, V 1-4, VI BASE 1-7
................................................ Pemphigus gairi
 Populus 361

48 ANT III 0.22-0.38 mm, but if less than 0.23 mm (small bursarius) then
ANT PT is short and stubby, less than 1.5 x its basal width . . . . . 49
- ANT III 0.15-0.25 mm, but if more than 0.22 mm then ANT PT is more
than 1.5 x its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
49 Primary rhinarium on ANT VI much larger than adjacent secondary
rhinaria and forming a broad band that extends over half-way around
antenna (Fig.87I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
- Primary rhinarium on ANT VI normal or irregular in shape but not
extending more than half-way around antenna . . . . . . . . . . . . . . . . . . 52
50 ANT VI BASE with 6-8 secondary rhinaria (Fig. 87I). (III 12-16, IV 3-5,
V 4-6, VI BASE 6-8) . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus passeki
- ANT VI BASE with 3-6 secondary rhinaria . . . . . . . . . . . . . . . . . . . . 51
51 ANT PT 0.03-0.04 mm. (Secondary rhinaria on I I I 10-15, I V 4-5, V 2-4,
VI BASE 3-6) . . . . . . . . . . . . . . . . . . . . . . . Pemphigus junctisensoriatus*
- ANT PT 0.02-0.03 mm. (Secondary rhinaria on I I I 12-16, I V 4-6, V 3-5,
VI BASE 3-6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus dorocola*
52 Tooth on inner side of ANT I I I often very close to base of segment,
usually without a rhinarium basad to it (Fig. 87J). (Secondary rhinaria on
III 8-13, IV 2-5, V 2-5, VI BASE 3-7) . . . . . Pemphigus populicaulis
- Tooth on inner side of ANT III almost always with 1-2 secondary rhinaria
basad to it . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
53 Secondary rhinaria on ANT I I I 8-17, IV 2-8, V 2-6, VI BASE 3-8. PT
usually less than 1.5 x basal width . . . . . . . . . . . . Pemphigus bursarius
- Secondary rhinaria on ANT III 9-13, IV 3-5, V 3-5, VI BASE 3-5. PT
usually more than 1.5 x basal width . . . . . . . . . Pemphigus mordvilkoi
54 Tooth on inner side of ANT III usually more than 0.07 mm from base
of segment, with always 1, and sometimes 2-3, secondary rhinaria basad
to it (Fig. 87K). (Secondary rhinaria on III 9-12, IV 2-4, V 2-4, VI BASE
3-5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus laurifoliae
- Tooth on inner side of ANT III usually less than 0.06 mm from base of
segment, with 0-1 secondary rhinaria basad to it . . . . . . . . . . . . . . . . 55
55 BL 2.1-2.5 mm. ANT V 0.8-1.0 x ANT IV, and more often than not
also bearing fewer secondary rhinaria than IV (examine several specimens;
secondary rhinaria distributed III 9-11, IV 0-5, V 0-4, VI 0-5) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populiramulorum
- BL 1.1-2.3 mm (only P. populiglobuli more than 2.0mm). ANT V
1.0-1.2 x ANT IV, and usually with at least as many sensoria as IV
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
56 ANT PT short, maximally about 0.03 mm long and usually less than
3 . 5 x its basal width. Tooth on ANT III usually without a rhinarium
362 Pouteria

basad to it (Fig. 87J). (Secondary rhinaria on III 8-13, IV 2-5, V 2-5, VI

BASE 3-7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populicaulis
- ANT PT usually longer than 0.03 mm, more than 1.5 x its basal width.
Tooth on ANT III often with a rhinarium basad to it . . . . . . . . . . . 57
56 Secondary rhinaria distributed III 5-13, IV 2-4, V 0-4, VI BASE 1-7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populiglobuli
- Secondary rhinaria distributed III 7-12, IV 1-5, V 2-5, VI BASE 3-7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus populivenae

POUTERIA ( = MALACANTHA) Sapotaceae

Pouteria alnifolia Aphis gossypii

P. ramiflora Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

PREMNA Verbenaceae

Premna odorata Mesotrichosiphum uichancoi

PROSOPIS Leguminosae

Prosopis stephaniana Aphis craccivora

PRUNUS Rosaceae

Prunus here includes Amygdalopsis, Amygdalus, Armeniaca, Cerasus, Lauro-

cerasus and Persica as subgenera. It is the primary host genus for most species
of the aphidine genera Brachycaudus, Hyalopterus, Rhopalosiphum, Myzus
and Tuberocephalus. Although some Prunus-feeding aphids are rather specific
in their choice of primary host, many colonize more than one species, and a
single key to aphids on Prunus of the world seems most useful.
 Prunus 363

Host Plant List

Prunus africana Aphis spiraecola
(Red Stinkwood, Bitter-Almond)
P. americana Aphis minima
(American Wild Plum, River Hyalopterus pruni
Plum)
P. amygdalus = P. dulcis
P. angustifolia Aphis minima
(Chickasaw Plum, Mountain
Cherry)
P. ansu Aphis gossypii
Myzus cerasi, umefoliae
P. armeniaca Asiphonaphis pruni
(Apricot) Brachycaudus cardui, persicae
Hyalopterus amygdali, pruni
Myzus mumecola, persicae
Pterochloroides persicae
Rhopalosiphum nymphaeae
(For a key to common aphids on Apricot, see B & E, 1984)
P. avium Brachycaudus helichrysi
(Sweet Cherry, Mazzard) Myzus cerasi, persicae
Nearctaphis bakeri
Rhopalosiphum nymphaeae
P. besseyi [Rhopalosiphum musae]
P. bucharica Brachycaudus pilosus
P. buergeriana [Myzus inuzakurae Shinji, 1930]
(Inuzakura, Shirozakura)
P. carolinaia Aphis spiraecola
(Carolina Laurelcherry) Myzus persicae
P. cerasifera (incl. var. pissardii) Aphis craccivora, fabae, spiraecola
(Cherry Plum) Brachycaudus helichrysi
Hyalopterus pruni
Myzus cerasi, lythri, persicae
Phorodon humuli
Pterochloroides persicae
P. cerasus [Betacallis prunicola]
(Sour Cherry, Morello Cherry) Brachycaudus cardui, [cerasicola],
helichrysi
Myzus cerasi, mushaensis, persicae,
yamatonis
Rhopalosiphum [insertum],
nymphaeae
364 Prunus

Tinocalloides montanus
Tuberocephalus higansakurae,
sakurae
P. cornuta Avicennina indica
Eumyzus pruni
Myzus cornutus
Pyrolachnus imbricatus
[Rhopalosiphum maidis]
Vesiculaphis pruni
P. davidiana Myzus persicae
P. divaricata Brachycaudus cardui, divaricatae,
[divaricatellus], helichrysi,
persicae, prunicola, schwartzi
Hyalopterus pruni
Phorodon humuli
Pterochloroides persicae
Rhopalosiphum nymphaeae
P. domestica Aphis longicauda, spiraecola
(Common or European Plum) Brachycaudus [almatinus], cardui,
divaricatae, helichrysi, persicae,
prunicola, semisubterraneus
Hyalopterus pruni
Hysteroneura setariae
Macrosiphum euphorbiae
Myzus ornatus, persicae
Nearctaphis baker i
Phorodon humuli
Pterochloroides persicae
Rhopalosiphum [musae],
nymphaeae, padi
(For a key to common aphids on Plum see B & E, 1984)
P. donarium = P. lannesiana
P. donarium var. sachalinensis = P. sachalinensis
P. donarium var, speciosa = P. jamasakura var. speciosa
P. donarium var. spontanea - P. serrulata var. spontanea
P. dulcis Aphis fabae, gossypii
(Almond) Brachycaudus amygdalinus,
helichrysi, persicae
[Capitophorus prunifoliae Shinji,
1924]
Hyalopterus amygdali
Myzus amygdalinus, persicae
 Prunus 365

Pterochloroides persicae
Rhopalosiphum nymphaeae
(For a key to common aphids on Almond see B & E, 1984)
P. emarginata Myzus cerasi, lythri
(Bitter Cherry)
P. erythrocarpa Aphidura bozhkoae
Brachycaudus pilosus
P. glandulosa Hyalopterus pruni
Myzus persicae, varians
Phorodon japonensis
Rhopalosiphum nymphaeae,
rufiabdominalis
Tuberocephalus momonis
P. grayana Myzus persicae
Tuberocephalus sp. (Sorin, 1993)
P. hortulana Hyalopterus pruni
P. humilis Tuberocephalus jinxiensis
P. incana Aphidura bozhkoae
P. insititia Brachycaudus cardui, helichrysi,
(Bullace, Damson) persicae, semisubterraneus
Brachycaudus (Appelia) prunicola
Hyalopterus pruni
Phorodon humuli
Rhopalosiphum nymphaeae
(For a key to common aphids on Damson see B & E, 1984)
P. itosakura var. ascendens Tuberocephalus sasakii
P. jamasakura (incl. var. speciosa) Myzus cerasi, mushaensis, persicae
(Yamasakura) Myzus (or Tuberocephalus) spp.
(Moritsu and Tokumotu, 1972)
Rhopalosiphum rufiabdominalis
Tuberocephalus sakurae, sasakii
P. japonica Phorodon japonensis
P. lannesiana Myzus mushaensis
(Oshima-zakura) Myzus (or Tuberocephalus) spp.
(Moritsu and Tokumotu, 1972)
Tuberocephalus sakurae, sasakii
P. laurocerasus Aulacorthum solani
(Cherry Laurel) Macrosiphum euphorbiae
Myzus persicae
P. lyonii Aphis spiraecola
366 Prunus

P. mahaleb Myzus amygdalinus, cerasi, lythri

Roepkea marchali, marchali ssp.
bathiaschvili
P. maximowiczii Prociphilus oriens
Tuberocephalus sakurae
P. melanocarpa = P. virginiana
P. microcarpa Brachycaudus pilosus
P. mume Brachycaudus helichrysi
(Ume, Japanese Apricot) Myzus cerasi ssp. umefoliae,
mumecola, persicae, yamatonis
Phorodon japonensis
Rhopalosiphum rufiabdorninalis
P. munsoniana Brachycaudus persicae
Myzus persicae
P. nakaii Rhopalosiphum rufiabdorninalis
P. nigra Hyalopterus pruni
(Canada Plum) Hysteroneura setariae
Myzus persicae
Rhopalosiphum cerasifoliae
P. padus (incl. var. pubescens) Eumyzus prunicola
(European Bird Cherry) Myzus asiaticus, padellus
Rhopalosiphum padi
[ Tuberocephalus momonis, sakurae]
P. pauciflora Tuberocephalus misakurae, sakurae
(Kara-misakura)
P. pendula = P. subhirtella
P. pennsylvanica Myzus cerasi
(Pin Cherry) Rhopalosiphum cerasifoliae, padi
P. persica Aphis spiraecola
(Peach) Brachycaudus amygdalinus,
helichrysi, persicae, persicaecola,
schwartzi
Hyalopterus amygdali, pruni
Hysteroneura setariae
Longistigma xizangensis
Macrosiphum euphorbiae
Myzus cerasi, [nicotianae], persicae,
varians
[Phorodon persifoliae Shinji,
1922c]
Pterochloroides persicae
 Prunus 367

Rhopalosiphum [momo Shinji

1922c], nymphaeae,
rufiabdominalis
Tinocalloides montanus
Tuberocephalus momonis
(For a key to common aphids on Peach see B & E, 1984)
P. pissardii = P. cerasifera var. pissardii
P. prostrata (?) Brachycaudus pilosus
P. pseudocerasus Myzus cerasi
Tuberocephalus higansakurae,
liaongensis, misakurae, sakurae
P. puddum Tinocalloides montanus
P. sachalinensis Prociphilus oriens
Tuberocephalus sakurae, sasakii
P. salicina Aphis gossypii
(Chinese or Japanese Plum) Brachycaudus persicae
Hyalopterus pruni
Myzus persicae
Phorodon japonensis
Pterochloroides persicae
Rhopalosiphum nymphaeae,
rufiabdominalis
P. sargentii = P. sachalinensis
P. serotina Aphis feminea
(Black Cherry) Asiphonaphis pruni
Hyalopterus pruni
Myzus cerasi, persicae
Rhopalosiphum padi
P. serrulata (incl. var. glabra, var. Myzus yamatonis
spontanea) Rhopalosiphum padi
(Japanese Flowering Cherry, Tuberocephalus artemisiae,
Oshima-zakura) higansakurae, momonis,
sakurae, sasakii
P. sieboldii Myzus cerasi
P. spinosa Brachycaudus cardui, divaricatae,
(Blackthorn or Sloe) helichrysi, prunicola,
semisubterraneus
Hyalopterus pruni
Phorodon humuli
Rhopalosiphum nymphaeae, padi
P. spinosissima Myzus persicae
368 Prunus

P. subcordata [Rhopalosiphum musae]

P. subhirtella (incl. var. pendula) Myzus persicae, yamatonis
(Ko-higanzakura, Rosebud Cherry) Tuberocephalus higansakurae
P. taiwaniana Myzus mushaensis
P. takenakae Tuberocephalus misakurae
P. tenella Myzus persicae
Rhopalosiphum padi
P. tianshanica Brachycaudus pilosus
P. tomemtosus Brachycaudus helichrysi, persicae
(Yusura-ume) Myzus persicae
Rhopalosiphum nymphaeae
Tuberocephalus momonis, sakurae
P. triflora Phorodon japonensis
P. triloba Hysteroneura setariae
Tuberocephalus sakurae
P. ulmifolia Aphis bozhkoae
Brachycaudus pilosus
P. umbellata Aphis minima
P. ursina Hyalopterus pruni
P. verrucosa Aphidura bozhkoae
Brachycaudus pilosus
P. virginiana (incl. var. demissa) Asiphonaphis pruni
(Choke Cherry) Hysteroneura setariae
Myzus cerasi
Rhopalosiphum cerasifoliae, padi
P. yedoensis Myzus cerasi, mushaensis, sakurae,
(Somei-yoshino) yamatonis
Rhopalosiphum rufiabdominalis
Tuberocephalus artemisiae, sakurae
Prunus spp. (unidentified) Aphis chetansapa
Aulacorthum magnoliae
[Myzus prunisuctus Zhang in
Zhang and Zhong, 1980a]
[Tuberocephalus tianmushaensis
Zhang in Zhang and Zhong,
1980a]

Key to aphids on Prunus spp.

Except where otherwise indicated, this key refers to apterous viviparae,
including fundatrices.
 Prunus 369

1 CAUDA knobbed, anal plate bilobed. SIPH short dark truncate cones,
shorter than their basal width, with a single hair on apical half. (All
viviparous females alate) . . . . . . . . . . . . . . . . . . . Tinocalloides montanus
- CAUDA rounded, triangular or finger-like, sometimes with a constriction
but not distinctly knobbed. Anal plate entire. SIPH of various forms, with
or without hairs, or absent. (Viviparae apterous or alate, rest of key
applies to apterae except where otherwise stated) . . . . . . . . . . . . . . . . . 2
2 SIPH as large pores on broad dark hairy cones . . . . . . . . . . . . . . . . . . 3
- SIPH tubular or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 Dorsum with a double row of large pigmented spinal tubercles. (Forewings
of alata maculate with a short blunt pterostigma) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterochloroides persicae
- Dorsum without spinal tubercles. (Forewings of alata not maculate,
pterostigma elongate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 EL less than 5.5 mm. (Alata with straight pterostigma, not reaching tip of
wing) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyrolachnus imbricatus
- BL more than 5.5 mm. (Alata with pterostigma curved around tip of wing)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma xizangensis*
5 ANT PT/BASE less than 0.3. SIPH absent, CAUDA broadly rounded.
(Forewing of aiata with an unbranched media) . . . . Prociphilus oriens
- ANT PT/BASE more than 0.8. SIPH present or absent, if absent then
CAUDA not broadly rounded. (Forewing of alata once- or twice-branched)
............................................................. 6

6 SIPH absent. Large lateral tubercles on thorax and ABD TERG 1-7
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiphonaphis pruni
- SIPH present. Lateral tubercles present or absent . . . . . . . . . . . . . . . . 7
7 Head capsule smooth, wrinkled or rugose, but not ornamented with
spicules or nodules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Head capsule with spiculose or nodulose ornamentation . . . . . . . . . . 30
8 SIPH small and thin, less than 0.7 x CAUDA, lacking an apical flange
(Figs 88A, B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- SIPH more than 0.7 x CAUDA, usually with an apical flange . . . . 10
9 SIPH very thin, 2.5-4.0 x their maximum diameter (Fig. 88A). (Alata
with total secondary rhinaria on both ANT III 41-74, on ANT IV 9-21)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalopterus pruni
- SIPH 1.6-2.5 x their maximum diameter (Fig. 88B). (Alata with total
secondary rhinaria on both ANT III 21-41, on ANT IV 0-10) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyalopterus amygdali
370 Prunus

Fig. 88. SIPH of aptera (in spring populations) of A, Hyalopterus pruni, B, H. amygdali, C, Avicennina
indica, D, Rhopalosiphum nymphaeae, E, Rh. cerasifoliae, F, Rh. padi.

10 SIPH dark, smooth and conspicuously inflated over more than half
length, with maximum diameter of swollen part 2 or more x minimum
diameter near base (Fig. 88C) . . . . . . . . . . . . . . . . . . . . Avicennina indica
- SIPH tapering, cylindrical, or if swollen then swelling is confined to distal
half, of maximum diameter less than 2 x minimum diameter of basal
half, and surface is rugose or imbricated . . . . . . . . . . . . . . . . . . . . . . . 11
11 Body spindle-shaped, SIPH long, 1.5-2.1 x the long finger-like CAUDA,
and with subapical zone of polygonal reticulation (Fig. 121C). ANT III
with 2-7 secondary rhinaria near base . . . . . Macrosiphum euphorbiae
- Body oval, if CAUDA is finger-like then SIPH are usually relatively
shorter, and always without polygonal reticulation. ANT III without
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 CAUDA tongue- or finger-like or tapering, usually more than 1.2 x its
basal width, but if shorter then with pointed apex . . . . . . . . . . . . . . . 13
- CAUDA bluntly conical, semicircular or broadly rounded, rarely up to
1.2 x its basal width, and never pointed . . . . . . . . . . . . . . . . . . . . . . . . 20
13 CAUDA long (more than 0.2 mm), conspicuously pale compared with the
black siphunculi, and bearing 4 hairs. Femora with contrasting black
distal and pale basal halves . . . . . . . . . . . . . . . . . . Hysteroneura setariae
 Prunus 371

- CAUDA usually dusky or dark, if pale then shorter than 0.2 mm and with
more than 4 hairs. Femora without contrasting black and pale sections
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14 Lateral tubercle on ABD TERG 7 situated posterioventrally to spiracle

(Fig. 42H), appearing in perfect dorsoventral view to be laterad to an
anterioposterior line drawn through spiracle. SIPH with basal diameter
usually at least 2 x external diameter of the small apical flange, without a
trace of a subapical swelling or constriction . . . . . . . . . . . . . . . . . . . . 15
- Lateral tubercle on ABD TERG 7 situated posteriodorsally to spiracle
(Fig. 42H), appearing in perfect dorsolateral view to be slightly mesiad to
an anterioposterior line drawn through spiracle. SIPH with basal diameter
less than 2 x external diameter of apical flange, and usually with a clear
subapical constriction, proximal to which there is some degree of
swelling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
15 CAUDA paler than siphunculi, w i t h 4-8 hairs . . . . . . . Aphis gossypii
- CAUDA black like siphunculi, and usually with 8-24 hairs . . . . . . . 16
16 CAUDA with a constriction just proximal to midpoint, and usually bear-
ing 8-12 hairs (Fig. 121T). Dorsal abdomen without any dark markings
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
- CAUDA without any constriction, and w i t h 11-24 hairs. Dorsal abdomen
at least with dark sclerites or bars on ABD TERG 7 and 8 (Fig. 121R)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis fabae
17 Tibiae evenly pigmented. SIPH more than 0.3 mm long (except fundatrix)
and swollen proximal to subapical constriction (Fig. 88D). (Fundatrix with
ANT PT/BASE more than 2.0) . . . . . . . . . Rhopalosiphum nymphaeae
- Tibiae mainly pale, dark only distally. SIPH less than 0.3mm long or,
if longer, without any discernible subapical swelling. (Fundatrix with
ANT PT/BASE 2.0 or less) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 ABD TERG 8 with 4-6 hairs. Hairs on ABD TERG 1-4 all more than
40 µm long with pointed apices . . . . . Rhopalosiphum rufiabdominalis
- ABD TERG 8 with 2-3 hairs. Hairs on ABD TERG 1-4 often with blunt
or expanded apices and less than 40 µm long . . . . . . . . . . . . . . . . . . . 19
19 SIPH 0.26-0.38 mm long (except fundatrix), cylindrical, with no dis-
cernible subapical swelling (Fig. 88E). ABD TERG 1-4 usually bearing
some spinopleural hairs with pointed apices, longer than basal diameter
of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum cerasifoliae
- SIPH 0.20-0.30 mm long, usually slightly swollen proximal to the
subapical constriction (Fig. 88F). ABD TERG 1-4 with all spinopleural
hairs short and blunt or slightly capitate, none exceeding basal diameter of
ANT III in length . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum padi
372 Prunus

20 Spiracular apertures small and oval, partly covered anteriorly by an oper-

cular plate. ABD TERG 1 and 7 always with well-developed lateral
tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- Spiracular apertures large and rounded, not partly covered by opercular
plates. ABD TERG 1 and 7 with lateral tubercles absent or only present
if there are well-developed tubercles on other tergites . . . . . . . . . . . . 23
21 Hairs on ANT III long and fine, mostly longer than basal diameter of seg-
ment. ABD TERG 2-4 (as well as 1 and 7) with prominent lateral
tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis feminea
- Hairs on ANT III shorter than basal diameter of segment. Lateral
tubercles not evident on ABD TERG 2-4 . . . . . . . . . . . . . . . . . . . . . . . 22
22 Very small aphid (BL less than 1.1 mm). SIPH less than 0.18 mm long.
Lateral tubercles on ABD TERG 1 and 7 elongate, much longer than their
basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis minima
- BL usually more than 1.1 mm, and SIPH more than 0.2 mm long. Lateral
tubercles on ABD TERG 1 and 7 not usually longer than their basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis chetansapa/longicauda
23 CAUDA very broadly rounded, less than 0.5 x its basal width. R IV + V
only about 0.8 x HT II and usually bearing 4 accessory hairs. ABD TERG
8 (or 7 and 8) with spinal tubercles . . . . . . Brachycaudus amygdalinus
- CAUDA more than 0.5 x its basal width. R IV+V at least 0.9 x HT II
and bearing at least 6 accessory hairs. ABD TERG 8 with or without spinal
tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 First tarsal segments each with 4 hairs (i.e. 2 sense pegs between a long
pair of hairs). Dorsal abdomen often extensively pigmented with broad
transverse bars, interrupted in the midline on at least ABD TERG 1 and
2 and separate from lateral sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
- First tarsal segments each with 2-3 hairs (0-1 sense pegs). Dorsal abdomen
pale or dark, but if dark there tends to be an uninterrupted shield, fused
across and between segments and sometimes incorporating lateral sclerites
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
25 Lateral abdominal tubercles (circular membranous areas) on ABD TERG
2-4 mostly greater than 18 µm in diameter, often as large as or larger than
spiracles, with adjacent lateral hairs less than half this diameter . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus schwartzi
- Lateral abdominal tubercles on ABD TERG 2-4, if present, mostly less
than 18 µm in diameter, with adjacent hairs more than half this diameter
and sometimes much longer . . . . . . . . . . . . . . . Brachycaudus prunicola
26 SIPH more than 0.1 x BL and clearly more than 2 x CAUDA. Anterior
part of mesosternum with a pair of separate mammariform tubercles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
 Prunus 373

- SIPH less than 0.1 x BL, about 2 x CAUDA or less. Mesosternum

without tubercles, or with tubercles fused across midline (divaricatae)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
27 R IV+V longer than 0.18mm. Dorsal abdomen variably pigmented,
sometimes pale. SIPH pale or dusky . . . . . . . . . . Brachycaudus cardui
- R IV+V shorter than 0.175mm. Dorsal abdomen always extensively
sclerotized, black. SIPH black . . . . . . . . . Brachycaudus persicae group
28 Dorsal abdomen pale (except fundatrix, which may have dark transverse
bars). CAUDA about as long as HT II or longer. Dorsal abdominal hairs
of variable length, often short . . . . . . . . . . . . . Brachycaudus helichrysi
- Dorsal abdomen dark, with long hairs. CAUDA shorter than HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 Hind femur pale. Pronotum with pleural and lateral tubercles. Lateral
tubercles usually present on all of ABD TERG 1-7. Dorsal abdominal
hairs thick and stiff, mostly with blunt apices . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus pilosus
- Hind femur mainly dark. Pronotum without tubercles. Lateral tubercles
absent or small, never on all of ABD TERG 1-7. Dorsal abdominal hairs
with finely pointed apices . . . . . . . . . . . . . . . . Brachycaudus divaricatae
30 Frons convex, with antennal tubercles completely undeveloped. SIPH
with closely-spaced transverse rows of small spinules or nodules (e.g.
Fig.42M . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
- Antennal tubercles at least partially developed. SIPH often heavily
imbricated, but not with closely-spaced rows of spinules . . . . . . . . . 32
31 CAUDA helmet-shaped, shorter than its basal width (Fig. 89A), and
about half as long as the short, dark siphunculi. Tergum mainly dark
(except fundatrices) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Roepkea marchali
- CAUDA usually a little longer than its basal width (Fig. 42D), and similar
in length to the short, pale siphunculi. Tergum pale or with variable dark
spots or patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis bakeri
32 Anterior part of mesosternum with a pair of spinal mammariform pro-
cesses, ornamented with spinules (Fig. 89B). Meso- and metathoracic
tergites and ABD TERG 1-7 almost completely dark, with a fine poly-
gonal pattern of reticulation. SIPH straight, tapering, dark, and almost
2 x CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphidura bozhkoae
- Mesosternum without spinal processes. Tergum either pale or dark, SIPH
variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33 ANT III with 1-2 secondary rhinaria near base. Antennal tubercles well
developed with inner faces approximately parallel in dorsal view . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
374 Prunus

- ANT III always without secondary rhinaria. If antennal tubercles are well
developed then the inner faces are usually gibbous in dorsal view or bear
projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
34 Head, legs and antennae mainly dark; femora basally pale but with distal
part black. SIPH clavate (Fig. 121E). CAUDA with a constriction . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
- Head, legs and antennae mainly pale. SIPH gradually tapering (Fig. 121K).
CAUDA without a constriction . . . . . . . . . . . . . . . . Aulacorthum solani
35 ANT I with a scabrous protruberance on inner side near apex. Antennal
tubercles (except in fundatrix) with a forwardly-pointed projection as long
as or longer than its basal width. SIPH long, pale and tapering . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
- ANT I with inner side scabrous or smooth but not protruberant. Antennal
tubercles without a finger-like projection. SIPH variable . . . . . . . . . 37
36 R IV+V more than 1.3 x HT II . . . . . . . . . . . . . . Phorodon japonensis
- R IV+V less than 1.3 x HT I I . . . . . . . . . . . . . . . . . . . Phorodon humuli
37 Antenna of aptera (= fundatrix) 4-segmented, w i t h ANT PT/BASE less
than 1. (Apterous fundatrigeniae not produced) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus sasakii
- Antenna of aptera (including f u n d a t r i x ) 5- or 6-segmented, with ANT
PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38 SIPH with one or more hairs (not always in fundatrix) . . . . . . . . . . 39
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
39 Dorsum pigmented (except fundatrix) . . . . . . . . . . . . . . . . . . . . . . . . . . 40
- Dorsum of cleared specimens pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
40 Femora spinulose . . . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus sakurae
- Femora without spinules . . . . . . . . . . . . . . Tuberocephalus liaoningensis
41 Tibiae scabrous along entire length . . . . . . . . Tuberocephalus momonis
- Tibiae with at least basal 0.75 of length smooth . . . . . . . . . . . . . . . . 42
42 SIPH about 2 x longer than basal width, shorter than ANT PT . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus jinxiensis*
- SIPH more than 3 x basal width, a little longer than ANT PT . . . 43
43 SIPH with 1-3 hairs in aptera (but not fundatrix), and 0-2 hairs in spring
migrant alata, which has secondary rhinaria distributed ANT III 44-60,
IV 14-22, V 1-5 . . . . . . . . . . . . . . . . . . . . . . . Tuberocephalus misakurae
- SIPH with 5-12 hairs in aptera (but not fundatrix), and 5-12 hairs in
spring migrant alata, which has secondary rhinaria distributed ANT III
15-25, IV 5-7, V 0-2 . . . . . . . . . . . . . . . . Tuberocephalus higansakurae
 Prunus 375

44 SIPH jet black distally with basal half contrastingly pale, the transition
between the two tones being rather abrupt. Antennae also contrastingly
banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus varians
- SIPH and antennal segments pale or dark, sometimes pale basally and
becoming darker distally, but not contrastingly two-toned . . . . . . . . 45
45 Longest hairs on ABD TERG 1-6 more than 20 µm long, with pointed
apices, 0.8 or more x basal diameter of ANT III. Hairs on ANT III with
blunt or pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
- Longest hairs on ABD TERG 1-6 less than 20 µm long, with blunt or
expanded apices, usually less than 0.8 x basal diameter of ANT III. Hairs
on ANT III always with blunt apices . . . . . . . . . . . . . . . . . . . . . . . . . . 51
46 Hairs on ABD TERG 1-6 t h i c k , rather spine-like, with distinctly tuber-
culate bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
- Hairs on ABD TERG 1-6 fine, w i t h o u t or with only very small tuberculate
bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
47 Dorsal cuticle strongly corrugated. R IV+V 0.8-0.9 x HT II. ANT
PT/BASE 2.20-2.42 (except in fundatrices) . . . . . . . . . Eumyzus pruni
- Dorsal cuticle not strongly corrugated. R IV+V 1.0-1.1 x HT I I . ANT
PT/BASE 2.60-3.22 (except in fundatrices) . . . Eumyzus prunicolus
48 SIPH black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus padellus
- SIPH pale or dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
49 SIPH less than 2.3 x CAUDA, which is more than 1.25 x its basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae (part)
- SIPH more than 2.3 x CAUDA, which is less than 1.25 x its basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
50 R IV+V 0.85-0.95 x HT II. Tibiae smooth. (Hind tibia of immatures
weakly spinulose) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus cornutus
- R IV+V 1.1-1.2 x HT II. Tibiae with spinules or imbrication at base.
(Hind tibiae of immatures smooth) . . . . . . . . . . . . . . . Myzus mumecola
51 SIPH dark, rather thin, narrow-based (length only about 6 x basal
diameter which is less than 0.06mm), with a slight subapical swelling
(Fig. 89C). Dorsal cuticle strongly corrugated . . . . . . Myzus yamatonis
- SIPH pale or dark, broad-based (length less than 6 x basal diameter,
and/or basal diameter more than 0.06mm). Dorsal cuticle variable,
sculptured or smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
52 Dorsum fully covered with a solid blackish sclerotic shield with warty
ornamentation. SIPH jet black, long and tapering (Fig. 89D) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus cerasi/umefoliae
376 Prunus

Fig. 89. A, CAUDA of aptera of Roepkea marchali; B, mesosternal processes of Aphidura bozhkoae;
SIPH of aptera of C, Myzus yamatonis, D, M. cerasi and E, M. mushaensis.

- Dorsum membranous or sclerotic, if sclerotic and rather dark then SIPH

are not jet black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
53 Dorsal abdomen with a distinctive pattern of dark intersegmental mark-
ings, comprising pleural and marginal dashes or spots between ABD
TERG 1-4 as well as paired spinal dashes in front of ABD TERG 1 and
between 4, 5 and 6 (Fig. 121J). BL 1.0-1.7 mm . . . . . . Myzus ornatus
- Dorsal abdomen without such a pattern of markings. BL 1.4-2.3 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
54 CAUDA with 4 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
- CAUDA with 5-9 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
55 SIPH less than 0.4 mm long (Fig. 89E), often wholly or distally dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus mushaensis
- SIPH more than 0.4 mm long, pale . . . . . . . . . . . Myzus amygdalinus*
56 R IV+V 0.83-0.92 x HT II. Dorsal cuticle sclerotic and warty or cor-
rugated, especially marginally . . . . . . . . . . . . . . . . . . Vesiculaphis pruni*
- R IV+V 0.92-1.40 x HT II. Dorsal cuticle membranous or lightly
sclerotized, smooth or wrinkled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
 Pseudotsuga 377

57 SIPH dusky to dark, 0.15-0.18 x BL. . . . . . . . . . . . . . . Myzus asiaticus

- SIPH pale or slightly dusky, 0.19 or more x BL . . . . . . . . . . . . . . . . 58
58 R IV+V 1.2-1.4 x HT II, with 2 accessory hairs. SIPH about
3 x CAUDA which bears 5-6 hairs . . . . . . . . . . . . . . . . . . . Myzus lythri
- R IV+V 0.9-1.2 x HT II, with 2-7 accessory hairs. SIPH less than
2 X CAUDA which bears 6-9 (usually 7) hairs . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae (part)

PSEUDOPANAX Araliaceae

Pseudopanax lessonii Myzus persicae

PSEUDOTSUGA Pinaceae

Pseudotsuga macrocarpa Cinara pseudotsugae

Essigella californica
P. menziesii ( = taxifolia) Adelges cooleyi
(Douglas-Fir) [Aphis fabae]
Cinara commatula, [dubia],
[osborni], pseudotaxifoliae,
pseudotsugae, splendens,
[taxifoliae], vagabunda
Essigella californica, wilsoni
Pachypappa rosettei
Prociphilus americanus (?)

Key to aphids on Pseudotsuga

1 Adult females with chitinous ovipositor . . . . . . . . . . . . . Adelges cooleyi
Adult females without chitinous ovipositor (viviparous) . . . . . . . . . . . 2
2 SIPH absent. Eyes of 3 facets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
SIPH present as pores or pigmented hairy cones. Eyes multifaceted
............................................................. 4
3 Antenna with hairs longer than basal diameter of ANT III. R IV+V with
about 20 accessory hairs. Head with wax pore plates. CAUDA dark
protruberant, with many long hairs . . . . . . . . . Prociphilus americanus
378 Psidium

Antenna with hairs shorter than basal diameter of ANT III. R IV+V
usually without any accessory hairs. Head without wax pore plates.
CAUDA not protruberant, with 2 short hairs . . . Pachypappa rosettei
4 Body elongate. Antenna 5-segmented. R V very short . . . . . . . . . . . . . 5
Body oval. Antenna 6-segmented. R V elongate . . . . . . . . . . . . . . . . . . 6
5 Second tarsal segments mostly dusky to dark. ANT PT at least
0.5 x diameter of rhinarium on ANT V. R IV+V 0.07-0.10 mm. Tarsal
claws with distinctly bifurcate apices (Fig. 73) . . . Essigella californica
Second tarsal segments mainly pale, sometimes dusky on distal third.
Primary rhinarium on ANT V exceptionally distal, so that length of PT
is greatly reduced, less than 0.5 x diameter of rhinarium. R IV+V
0.04-0.07 mm. Tarsal claws with apices double but not distinctly bifurcate
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Essigella wilsoni
6 SIPH pores on broad pigmented conical bases 0.4mm or more in
diameter. (Antennal hairs numerous, 80 µm or more long; hind tibial hairs
100-150 µm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara commatula*
SIPH cones very small, with pigmented area less than 0.2 mm in diameter
............................................................. 7
7 All tibiae pale except at apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
At least hind tibia with distal third or more dark . . . . . . . . . . . . . . . . 9
8 Longest hairs on dorsal side of hind tibia less than 65 µm long . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara splendens
Longest hairs on dorsal side of hind tibia 90-160 µm long . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pseudotsugae
9 Longest abdominal hairs more than 100 µm long . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara vagabunda*
Longest abdominal hairs less than 100µm long . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara pseudotaxifoliae

PSIDIUM Guava Myrtaceae

Psidium guajava Aphis craccivora, gossypii,

spiraecola
Greenidea decaspermi, neoficicola
Greenidea (Trichosiphum)
formosana
Myzus ornatus, persicae
Toxoptera aurantii
 Píerocarya 379

Key to aphids on Psidium

1 SIPH without hairs . . . . . . . . . . use key to polyphagous aphids, p. 532
SIPH with numerous long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH with reticulation only at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea (Trichosiphum) formosana
SIPH with a pattern of pale reticulation extending over almost their entire
length ....................................................... 3
3 SIPH black, paler at apices, and longer than hind tibiae . . . . . . . . . . . .
........................................... Greenidea decaspermi
SIPH dusky yellow, darker at apices, not longer than hind tibiae . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea neoficicola

PSYCHOTRIA Rubiaceae

Psychotria grandis Aphis gossypii

Aulacorthum solani
(Use key to polyphagous aphids, p. 532)

PTEROCARPUS Leguminosae

Pterocarpus soyauxii Aphis craccivora

(African Padouk, Camwood) Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

PTEROCARYA Wing-nuts Juglandaceae

Pterocarya fraxinifolia [Chromaphis californica]

(Caucasian Wing-nut) [Eucarazzia caucasica Aizenberg,
1956]
[Monelliopsis bisetosa]
(all probably vagrants)
380 Pterospermum

Pt. rhoifolia Kurisakia [ailanthi ssp.

(Japanese Wing-nut) sawagarumii], onigurumii
Myzus persicae
Prociphilus oriens
Pt. stenoptera [Aphis fabae]
(Chinese Wing-nut) Dasyaphis rhusae
Kurisakia onigurumii (?)
[Tinocallis insularis]

Key to aphids on Pterocarya spp.

1 SIPH tubular. ANT PT/BASE much more than 1 . . . . . . . . . . . . . . . 2
SIPH broadly conical, pore-like or absent. ANT PT/BASE less than 1
............................................................. 3
2 SIPH much inflated on distal half with maximum diameter of swollen part
more than 2 x minimum diameter of 'stem' . . . . Eucarazzia caucasica
SIPH not or only slightly clavate, with maximum diameter much less than
2 x minimum diameter . . . . . go to key to polyphagous aphids, p. 532
3 Aptera with 3-segmented antennae, and body often covered with long
hair-bearing processes. CAUDA knobbed, anal plate divided deeply into
2 triangular lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dasyaphis rhusae
Aptera with 5-segmented antennae, body without long processes. CAUDA
rounded, anal plate entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 SIPH as large pores on shallow cones, ringed with long hairs. Alata with
5-segmented antennae and media of forewing once-branched . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kurusakia onigurumii
SIPH absent. Alata with 6-segmented antennae and media unbranched
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens

PTEROSPERMUM Sterculiaceae

Pterospermum spp. Cervaphis schouteniae

Eutrichosiphum subinoyi

Key to apterae on Pterospermum

Body with very long, branched, hair-bearing marginal processes. SIPH
cylindrical with sparse hairs . . . . . . . . . . . . . . . . . Cervaphis schouteniae
Body without long branched marginal processes. SIPH swollen, with
numerous long hairs . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum subinoyi
 Pyrus 381

PTEROSTYRAX Styracaceae

Pterostyrax corymbosum Myzus persicae

PYRUS Pears Rosaceae

A key to the common aphids on cultivated pear (P. communis) is provided

by B & E (1984). A revised key, extended to include all aphids recorded from
the genus Pyrus, is given here. Aphids on crab-apples, often included in
Pyrus, are treated under Malus.

Host Plant List

Pyrus amygdaliformis Dysaphis pyri
P. boissieriana Dysaphis reaumuri
P. bondoensis Sappaphis piri
P. communis Anuraphis catonii, farfarae,
(Common Pear) pyrilaseri, subterranea
Aphanostigma piri
Aphis craccivora, gossypii, pomi,
spiraecola
Brachycaudus [almatinus], cardui,
helichrysi, persicae
Dysaphis [multisetosa], plantaginea,
pyri, reaumuri, tschildarensis
Eriosoma americana, flavum,
lanigerum, lanuginosum,
pyricola
[Eutrichosiphum pyri]
Fimbriaphis gentneri
Longistigma caryae, xizangensis
Melanaphis pahanensis, pyraria
Myzus persicae
Nearctaphis bakeri
Nippolachnus bengalensis, pyri
Ovatus crataegarius, insitus
Prociphilus kuwanai
Pterochloroides persicae
Pyrolachnus pyri
382 Pyrus

Rhopalosiphum insertum
Sappaphis piri
Schizaphis punjabipyri, pyri
Toxoptera aurantii, citricidus,
odinae
P. granulosa Nippolachnus bengalensis
P. korshinskyi Dysaphis pyraria, reaumuri
P. kumaoni Melanaphis pahanensis
P. lanata [Dysaphis pavlovskyana]
P. montana Aphis spiraecola
[Brachysiphoniella montana]
Sappaphis piri
P. nivalis Dysaphis pyri
P. pashia Dysaphis sharmai
Melanaphis pahanensis
Nippolachnus bengalensis
Pyrolachnus pyri
P. pulcherrima Aphis gossypii
P. pyrifolia (incl. var. culta) Aphanostigma iaksuiense
(Dolbae, Chinese or Japanese Pear) Aphis gossypii, pomi, spiraecola
Aulacorthum magnoliae
Brachycaudus helichrysi
Longistigma caryae
Melanaphis siphonella
Myzus persicae
Nippolachnus piri
Ovatus crataegarius
Prociphilus kuwanai, oriens
Rhopalosiphum insertum,
nymphaeae, rufiabdominalis
Sappaphis piri
Schizaphis piricola
Toxoptera odinae
P. salicifolia Dysaphis reaumuri
P. syriaca Aphanostigma piri
Dysaphis reaumuri
P. ussuriensis Dysaphis pyri
Prociphilus kuwanai
Sappaphis piri
P. vestita Eurnyzus eastopi
 Pyrus 383

Pyrus spp. Aphis chetansapa

[Dysaphis affinis]
Sappaphis dipirivora, sinipiricola
Sinomegoura pyri

Key to aphids on Pyrus (apterous viviparae, excluding fundatrices, except

where otherwise stated)
1 Antenna 3-segmented. Body very small (BL 0.8-1.0 mm), pyriform,
broadest anteriorly. All females oviparous . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphanostigma iaksuiense/piri
- Antenna 4- to 6-segmented. Body usually longer than 1 mm, not broadest
anteriorly. Parthenogenetic females viviparous . . . . . . . . . . . . . . . . . . . 2
2 ANT PT/BASE less than 0.8; if more than 0.6 then PT bears some long
fine hairs. CAUDA broadly rounded, usually much shorter than width at
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- ANT PT/BASE more than 0.6, and PT without any long fine hairs.
CAUDA semicircular, helmet-shaped, tongue- or finger-shaped, about as
long as its basal width or longer, never much shorter . . . . . . . . . . . . 15
3 Conspicuous wax pore plates present. SIPH either absent or present as
pores with a surrounding ring of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Wax pore plates absent. SIPH as large pores on broad cones with
numerous hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
4 SIPH present as large pores with surrounding hairs. Wax pore plates com-
prising rings or groups of circular or polygonal facets enclosing one or
more clear central areas (on stems or roots) . . . . . . . . . . . . . . . . . . . . . 5
- SIPH absent or inconspicuous. Wax pore plates honeycomb-like without
any clear central areas (in leaf-nest galls) . . . . . . . . . . . . . . . . . . . . . . . . 9
5 All wax pore plates consisting of a ring of facets around a single, more-or-
less circular central clear area (e.g. Fig. 109A, p. 485) . . . . . . . . . . . . . 6
- Largest wax pore plates each consisting of a group of facets enclosing
either a very narrow or subdivided central clear area or several clear
areas (Fig.60C, p.239) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
6 BL 2.0-2.7 mm. R IV+V 0.22-0.25 mm. (CAUDA with 2-4 hairs) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanuginosum
- BL 1.0-1.9mm. R IV+V 0.10-0.18 mm . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 BL 1.0-1.3 mm. R IV+V 0.10-0.12 mm. CAUDA with 2-4 hairs . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma flavum
- BL 1.3-1.9mm. R IV+V 0.12-0.18mm. CAUDA with 5 or more hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma pyricola
384 Pyrus

8 ANT III similar in length to R IV+V. Wax pore plates with narrow, mostly
interconnecting central areas . . . . . . . . . . . . . . . . . . Eriosoma lanigerum
- ANT III less than 0.5 x R IV+V. Wax pore plates with separate, circular
or polygonal clear central areas . . . . . . . . . . . . . . Eriosoma americanum
9 BL of spring migrant alata from leaf-nest gall 4mm or more. R IV+V
more than 0.2 mm long and bearing 7 or more accessory hairs . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
- BL of spring migrant alata less than 4 mm. R IV+V less than 0.16 mm long
and bearing 2-5 accessory hairs . . . . . . . . . . . . . . . Prociphilus kuwanai
10 Body elongate, 2 or more x longer than wide. Antennae and legs pale
except for black tibial apices and hind tarsi. (Alata with a black quadrate
dorsal abdominal patch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Body broadly oval, less than 2 x longer than wide. Antennae and legs
mainly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 ANT PT/BASE 0.25-0.35, without any long fine hairs distal to primary
rhinarium. R I V + V 0.14-0.18 mm long, 0.62-0.75 x HT II, and bearing
9-16 accessory hairs . . . . . . . . . . . . . . . . . . . . . Nippolachnus bengalensis
- ANT PT/BASE 0.45-0.75, with some long fine hairs distal to primary
rhinarium. R IV+V 0.17-0.22 mm long, 0.7-1.0 x HT II, and bearing
more than 20 accessory hairs . . . . . . . . . . . . . . . . . . . . Nippolachnus piri
12 Abdomen with paired, pigmented spinal tubercles on each tergite. Alata
with maculate forewings and a short pterostigma . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterochloroides persicae
- Abdomen without spinal tubercles. Forewings of alata not maculate, with
a long pterostigma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 HT I (measured on ventral side) less than 2 x its basal diameter, less than
0.35 x HT II. Forewing of alata with pterostigma almost straight, not
extending around tip of wing . . . . . . . . . . . . . . . . . . . . Pyrolachnus pyri
- HT I more than 2 x its basal diameter, more than 0.35 x HT II. Forewing
of alata with pterostigma curved distally, extending around wing-tip
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 HT II 1.8-2.3 x HT I . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- HT II about 2.6 x HT I . . . . . . . . . . . . . . . . . Longistigma xizangensis*
15 SIPH shorter than CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- SIPH as long as or longer than CAUDA . . . . . . . . . . . . . . . . . . . . . . . 20
16 Abdomen without any dorsal markings . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Abdomen with dark dorsal markings . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 Stridulatory ridges present on abdominal sternites 5 and 6. CAUDA dark,
rounded at apex (Fig. 121O) . . . . . . . . . . . . . . . . . . . . . Toxoptera odinae
 Pyrus 385

- No stridulatory ridges. CAUDA pale or dusky, long and tapering ...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachysiphoniella montana
18 SIPH very short, conical, shorter than basal width and about 0.2 x
CAUDA. Dorsal abdomen with interrupted markings . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melanaphis siphonella
- SIPH more than 0.7 x CAUDA. Dorsal abdomen with an almost solid
dark shield, or with dark transverse sclerites . . . . . . . . . . . . . . . . . . . . 19
19 Dorsal abdomen usually with an almost solid dark sclerotic shield. Hairs
on antenna and dorsal body minute, less than 0.5 x diameter of ANT III.
SIPH about 2 x basal width . . . . . . . . . . . . . . . . . . . Melanaphis pyraria
- Dorsal abdomen with separate transverse sclerites. Hairs on antennae and
dorsal body long and fine, much longer than diameter of ANT III. SIPH
less than 1.5 x basal width . . . . . . . . . . . . . . . . . Melanaphis pahanensis
20 SIPH with closely-spaced rows of densely-packed spinules or nodules
(e.g. Fig. 42M) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- SIPH often imbricated but without densely-packed spinules or nodules
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
21 Abdomen of aptera and alata without spinal tubercles. Alata with dark
dorsal patch centred on ABD TERG 3-6. Marginal tubercles if present
somewhat protuberant, and always absent from ABD TERG 7 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis bakeri
- Abdomen of alata with rather flat round spinal and marginal tubercles
on most tergites (present or absent on 7), and with dark patch centred
on ABD TERG 4-6 or 5-6 (ABD TERG 3 unpigmented). (Forming
pseudogalls inhabited by fundatrices and alate spring migrants only)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 Fundatrix with ANT PT/BASE greater than 1, and with well-developed
marginal sclerites that are mostly more than 2 x diameter of any tubercles
present. Alata with spinal and marginal tubercles usually on ABD TERG
1-7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
- Fundatrix with ANT PT/BASE less than 1, and with marginal tubercles
on small sclerites mostly less than 2 x diameter of tubercles. Alata with
spinal and marginal tubercles on ABD TERG 1-5 . . . . . . . . . . . . . . . 24
23 Fundatrix without spinal or marginal tubercles. SIPH of alata with 18-27
rows of spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anuraphis catonii
- Fundatrix with large marginal tubercles on ABD TERG 1-7, and often
also with spinal tubercles. SIPH of alata with 25-35 rows of spinules
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anuraphis subterranea
24 Fundatrix usually with a marginal tubercle on ABD TERG 6. Alata with
ANT PT/BASE 3.5-4.5 . . . . . . . . . . . . . . . . . . . . . . . Anuraphis farfarae
386 Pyrus

- Fundatrix without a marginal tubercle on ABD TERG 6. Alata with ANT

PT/BASE more than 5.0 . . . . . . . . . . . . . . . . . . . . . Anuraphis pyrilaseri
25 Dorsal cuticle sclerotic and strongly corrugated. R IV+V 1.68-1.90 x HT
II and bearing 2 accessory hairs . . . . . . . . . . . . . . . . . . Eumyzus eastopi
- Dorsal cuticle not strongly corrugated. R IV+V either less than 1.5 x HT
II or with more than 2 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 CAUDA tongue- or finger-shaped, rounded or pointed at apex, 1.2 or
more x longer than its basal width in dorsal view . . . . . . . . . . . . . . . 27
- CAUDA helmet-shaped, bluntly conical or rounded in dorsal view, not
clearly longer than its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
27 Head capsule spinulose or nodulose. Antennal tubercles well developed
with inner faces spinulose or scabrous and parallel or convergent in dorsal
view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
- Head capsule smooth. Antennal tubercles either small or undeveloped or
with inner faces smooth and divergent in dorsal view . . . . . . . . . . . . 31
28 Head, antennae and legs mainly dark. Inner faces of antennal tubercles
approximately parallel, and spinulose (e.g. Fig. 42J) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
- Head, legs and antennae mainly pale. Inner faces of antennal tubercles
convergent and scabrous (e.g. Fig. 42I) . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 SIPH moderately swollen on distal half. Alata with dark dorsal abdominal
patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
- SIPH cylindrical or tapering on distal half. Alata without dorsal
abdominal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 Spring migrant alata with 60-83 secondary rhinaria on ANT III, 36-52
on ANT IV and 13-22 on ANT V . . . . . . . . . . . . . . . . . . Ovatus insitus
- Spring migrant alata with 22-49 secondary rhinaria on ANT III, 5-20 on
IV and 0-10 on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus crataegarius
31 SIPH and CAUDA pale, mainly concolorous with dorsal cuticle. Anten-
nal tubercles well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
- SIPH, and usually CAUDA also, mainly darker than dorsal abdomen
(except when tergum is black). Antennal tubercles poorly developed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
32 ANT III with a single rhinarium near base and bearing minute hairs less
than 0.3 x basal diameter of segment. CAUDA with a slight mid-way
constriction and blunt apex, and usually bearing 6 hairs . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fimbriaphis gentneri
- ANT III with 2 rhinaria near base and hairs 0.5-0.7 x basal diameter of
segment. CAUDA long and pointed, with 11-12 hairs . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinomegoura pyri*
 Pyrus 387

33 Marginal tubercles on ABD TERG 7 placed just posteriad to spiracle and

slightly dorsad of an anterioposterior line drawn through spiracle; these
tubercles not usually of larger basal diameter than the spiracular opening
(as in Fig. 42H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
- Marginal tubercles on ABD TERG 7 placed posterioventrad to spiracle
and usually larger than the spiracular opening (Fig. 42H) . . . . . . . . . 39
34 SIPH tapering gradually from base to apex, not constricted subapically
and with flange only moderately developed . . . . . . . . . . . . . . . . . . . . . 35
- SIPH approximately cylindrical for most of length, slightly swollen
subapically, and constricted just proximad to the well-developed flange
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
35 Anterior part of head dark. SIPH uniformly black. Alata without any
secondary rhinaria on ANT V (III with 8-11, IV with 1-5) . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizaphis punjabipyri
- Head wholly pale or dusky. SIPH either uniformly dark or dusky with
black apices. Alata with 1-6 secondary rhinaria on ANT V (rarely 0 on
one side) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 Antennae shorter than body, reaching to base of siphunculi. SIPH
uniformly dark. Alata with secondary rhinaria distributed ANT III 14-19,
IV 5-9, V 0-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizaphis pyri
- Antennae about as long as body. SIPH pale or dusky, black only at apices.
Alatae with secondary rhinaria distributed ANT III 24-32, IV 14-16,
V 4-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizaphis piricola
37 Tibiae evenly pigmented. SIPH more than 0.3 mm long (Fig. 88D) and
clavate proximad to subapical constriction (except fundatrix). (Fundatrix
with ANT PT/BASE more than 2.0) . . . . Rhopalosiphum nymphaeae
- Tibiae mainly pale, dark distally. SIPH less than 0.3 mm long, with only
a slight swelling proximad to subapical constriction . . . . . . . . . . . . . . 38
38 ABD TERG 8 with 4-6 hairs. Hairs on ABD TERG 1-4 all more than
40 µm long with pointed apices. (Fundatrix with ANT PT/BASE equal to
or less than 2.0) . . . . . . . . . . . . . . . . . . . Rhopalosiphum rufiabdominalis
- ABD TERG 8 with 2 (-3) hairs. Hairs on ABD TERG 1-4 often with blunt
apices and less than 40 µm long. (Fundatrices with ANT PT/BASE more
than 2.0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphum insertum
39 Stridulatory ridges present on ventral abdomen (Fig. 122) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii, citricidus
(for separation see key to polyphagous aphids, p. 532)
- Stridulatory ridges absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . Aphis craccivora, eugeniae, fabae, gossypii, pomi, spiraecola
(for separation see couplets 21-25 of key to aphids on Malus)
388 Pyrus

40 First tarsal segments all with only 2 hairs. Marginal tubercles well
developed on ABD TERG 1 and 7 . . . . . . . . . . . . . . . Aphis chetansapa
- First tarsal segments with 3 hairs at least on fore and mid-legs. Marginal
tubercles if present then on ABD TERG 2-5 (or 2-6) as well as on 1
and 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41 Antenna with numerous long very fine hairs, the longest more than
2 x basal diameter of ANT III. (Alata with separate transverse bars on
dorsal abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
- Longest hairs on ANT III less than 1.8 x basal diameter of segment.
SIPH without hairs. (Alata with a more-or-less solid dark patch on dorsal
abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
42 SIPH 2 or more x basal width, and sometimes bearing 1 or more hairs.
CAUDA rounded, much shorter than its basal width . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sappaphis piri
- SIPH less than 1.5 x basal width, without hairs. CAUDA bluntly
pointed, about as long as its basal width . . . . . Melanaphis pahanensis
43 Spiracular openings large and rounded . . . . . . . . . . . . . . . . . . . . . . . . . 44
- Spiracular openings small and reniform . . . . . . . . . . . . . . . . . . . . . . . . 46
44 SIPH less than 2 x CAUDA. Dorsal abdomen dusky or pale. Meso-
sternum without mammariform processes . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus almatinus/helichrysi
- SIPH more than 2 x CAUDA. Dorsal abdomen usually dark. Anterior
part of mesosternum with a pair of mammariform processes . . . . . 45
45 R IV+V more than 0.17 mm, more than 0.6 x SIPH. Dorsal abdomen and
SIPH pale or dark. (Alata with secondary rhinaria only on ANT III)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus cardui
- R IV+V less than 0.17mm, about 0.5 x SIPH or less. Dorsal abdomen
always extensively sclerotized, black, and SIPH black. (Alata with secon-
dary rhinaria on ANT III and IV, or III, IV and V) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus persicae
46 Antennae at least as long as distance from frons to SIPH bases (in alata
about as long as body). SIPH more than 0.3 mm long, black . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis plantaginea
- Antennae shorter than distance from frons to SIPH bases (in alata less
than BL). SIPH pale or dark, less than 0.3 mm long . . . . . . . . . . . . 47
47 R IV+V 0.18-0.21 mm, 1.4-1.5 x HT II. Marginal tubercles present on
ABD TERG 1-5 and 7. Spring migrant alata (s.m.a.) with 13-22 second-
ary rhinaria on ANT I I I , 2-5 on IV, 0 on V . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis tschildarensis
 Quercus 389

- R IV+V 0.12-0.17 mm, 0.8-1.3 x HT II. Marginal tubercles present on

ABD TERG 1-5 or 1-7. S.m.a. where known with 23-64 secondary
rhinaria on ANT III, 2-27 on IV and 0-7 on V . . . . . . . . . . . . . . . . 48
48 SIPH 0.10-0.13 mm, 0.8-1.0 x R IV+V. S.m.a. with strongly protu-
berant secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . Dysaphis sharmai
- SIPH 0.16-0.29 mm, 1.2-2.0 x R IV+V. S.m.a. (where known) with
secondary rhinaria weakly protuberant . . . . . . . . . . . . . . . . . . . . . . . . . 49
49 Marginal tubercles present on ABD TERG 1-7 (sometimes missing from
one side on 6); slightly convex, the largest 55-60 µm in diameter. Longest
hairs on ANT III 12-18 µm, 0.5-0.8 x basal diameter of segment . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis pyraria*
- Marginal tubercles usually present only on ABD TERG 1-5 (very rarely
on 7 on one side); hemispherical, papilliform or conical, the largest
21-47 µm in basal diameter. Longest hairs on ANT III 13-50 µm,
0.5-1.8 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
50 SIPH black, 3.0-4.1 x their diameter at midpoint. Marginal tubercles
hemispherical. S.m.a. with 23-36 secondary rhinaria on ANT III, 2-10
on IV and 0-1 on V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis pyri
- SIPH pale, sometimes dusky at apices, 4.1-5.0 x their diameter at mid-
point. Marginal tubercles papilliform or conical. S.m.a. with 32-64
secondary rhinaria on ANT I I I , 8-27 on IV and 0-7 on V . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis reaumuri

QUALEA Vochysiaceae

Qualea multiflora Aphis gossypii, spiraecola

(Use key to polyphagous aphids, p. 532)

QUERCUS Oaks Fagaceae

About 225 aphid species in more than 40 genera feed on oaks. Some species,
notably of Hormaphidinae, seem restricted to subgenus Cyclobalanopsis, but
within the main subgenus Euquercus there is no clear pattern of host
specificity relating to the sections of the subgenus; e.g. the North American
aphid species complexes within Myzocallis, subgenera Neomyzocallis and
Lineomyzocallis, are thought to be mainly associated with red oaks (sect.
Erythrobalanus), yet many species in these groups are sometimes recorded
390 Quercus

from American white oaks (sect. Leucobalanus). There is therefore no alter-

native to keying all Quercus aphids together. A single key would be too long
and cumbersome, so after the host plant list there is a master key to the aphid
subfamilies associated with Quercus followed by keys to the species in each
subfamily (Drepanosiphinae being divided into two keys).

Host Plant List

Q. acuta Allotrichosiphum kashicola
(Japanese Evergreen Oak, Eutrichosiphum davidi, pasaniae
Aka-gashi) Greenidea kuwanai
Indonipponaphis fulvicola
Lachnus tropicalis
Nipponaphis distyliicola
Xenothoracaphis kashifolia
Q. acutissima Cervaphis quercus
(Japanese Chestnut Oak, Kunugi) Diphyllaphis konarae, quercus
Eutrichosiphum tattakanum
Greenidea kuwanai, nipponica
Kurisakia querciphila
Lachnus takahashii, tropicalis
Myzocallis kuricola
Nipponaphis distyliicola
Phylloxera kunugi
Stomaphis japonica
Tuberculatus acuminatus. capitatus,
fangi, fuscotuberculatus,
grisipunctatus, indicus, japonicus
ssp. radisectuae, konaracola
ssp. ganzuensis and ssp.
hangzhouensis, [kunugi Shinji,
1924], pappus, quercicola,
stigmatus, yokoyamai
Q. aegilops Lachnus sp., swirskii
Myzocallis castanicola, komareki
Thelaxes dryophila, suberi,
valtadorosi
Tuberculatus eggleri, pallidus
Q. agrifolia Atarsaphis agrifoliae
(California Live Oak) Myzocallis agrifolicola, walshii
Neosymydobius agrifoliae
Stegophylla essigi
Tuberculatus maureri, pallidus
Q. alba [Chaitophorus quercifoliae Fitch,
(American White Oak) 1851]
 Quercus 391

Hoplochaitophorus quercicola
Myzocallis bella, discolor, mimica,
punctata, walshii
Neosymydobius albasiphus,
mimicus
Phylloxera querceti, rileyi, ?stellata
Stegophylla querci
Stomaphis quercus
Thelaxes californica
Tuberculatus pallidus,
tuberculatus
Q. aliena Diphyllaphis quercus
(Galcham, Korean Oak) Greenidea nipponica
Tuberculatus capitatus, indicus,
querciformosanus, stigmatus,
yokoyamai
Q. arizonica Tuberculatus pallidus
Q. bicolor Hoplochaitophorus spiniferus
(Swamp White Oak) Myzocallis discolor, multisetis,
punctatus, walshii
Phylloxera rileyi
Thelaxes dryophila
Q. boisieri = var. of Q. infectoria

Q. borealis = Q. rubra
Q. calliprinos Hoplocallis microsiphon, [pictus]
(Prickly-leaved Evergreen Oak) Lachnus roboris
Myzocallis cocciferina, ?tauricus
Phylloxera quercus
Siculaphis vittoriensis
Thelaxes suberi
Tuberculatus maculipennis,
pallescens
Q. canariensis Myzocallis mediterraneus
(Mirbeck's Oak) Tuberculatus africanus, annulatus
Q. castanaefolia Myzocallis boerneri, castanicola
(Boland Masu, Persian Oak) Thelaxes suberi
Tuberculatus annulatus, pallidus
Q. castanea Myzocallis pepperi ssp. iturbide
(Encino, Mexican Oak) Stegophylla mugnozae
Q. catesbei = Q. laevis
392 Quercus

Q. centralis Neosymydobius butzei var.

papillata
Q. cerris Diphyllaphis mordvilkoi
(European Turkey Oak) Hoplocallis microsiphon, pictus,
ruperti
Lachnus roboris
Myzocallis bella, boerneri,
castanicola, komareki
Phylloxera quercina
Stomaphis quercus
Thelaxes suberi
Tuberculatus borealis, eggleri
Q. chrysolepis Neosymydobius agrifoliae,
(Canyon Live Oak) chrysolepis
Tuberculatus chrysolepidis
Q. cinerea Myzocallis longiunguis, multisetis,
(Bluejack Oak) spinosa
Neosymydobius luteus
Q. coccifera Lachnus roboris
(Holly Oak, Kermes Oak) Myzocallis castanicola, cocciferina,
komareki
Phylloxera quercus
Siculaphis vittoriensis
Thelaxes suberi, valladorosi
Q. coccinea Myzocallis bella, discolor,
(Scarlet Oak) longiunguis, melanocera,
multisetis, spinosa
Q. conferta = Q. frainetto
Q. crassipes Mexicallis spinifer
Myzocallis pepperi ssp. iturbide,
tenochca
Stegophylla mugnozae
Q. crispula Greenidea kuwanai
(Japanese White Oak) Lachnus tropicalis
Neothoracaphis yanonis
Nipponaphis distyliicola
Tuberculatus [naganoe Shinji,
1941], yokoyamai
Q. dealbata Allotrichosiphum assamense
Eutrichosiphum dubium. flavum,
khasyanum, rameshi, russellae
Indonipponaphis tuberculata
 Quercus 393

Lachnus acutihirsutus, tropicalis

Myzocallis polychaeta
Serratocallis takahashii
Q. dentata (incl. var. grandifolia) Diphyllaphis konarae
(Daimyo Oak, Kashiwa) Eutrichosiphum shiicola
Greenidea kuwanae, nipponica
Lachnus tropicalis
Metanipponaphis cuspidatae
Neothoracaphis yanonis
Phylloxera glabra, querceti
Tuberculatus annulatus, capitatus,
higuchii, indicus, japonicus,
kashiwae, [naganoe Shinji,
1941], neglectus, paiki, quer-
cicola, querciformosanus,
stigmatus, yokoyamai
Q. dilatata Eutrichosiphum assamense
(Moru Oak, Himalaya Oak) Globulicaudaphis pakistanica
[Myzocallis floribundi Verma,
1965]
Q. douglasii Neosymydobius agrifoliae
(Blue Oak) Phylloxera stanfordiana
Stegophylla essigi
Tuberculatus annulatus, quercifolii
Q. dumosa Stegophylla essigi
Tuberculatus pallidus, passalus
Q. durandii Stegophylla quercifoliae
Q. durifolia Myzocallis durangoensis
Q. emoryi Tuberculatus pallidus
Q. engelmannii Myzocallis punctatus
Neosymydobius chrysolepis
Tuberculatus pallidus
Q. engleriana Myzocallis castanicola
Q. fabri Tuberculatus capitatus, japonicus
ssp. radisectuae
Q. faginea Thelaxes suberi
Q. falcata (incl. var. pagodifolia) Myzocallis longiunguis, multisetis
(Southern Red Oak, Spanish Oak)
Q. farnetto = Q. frainetto
Q. fenestrata Eutrichosiphum assamense
394 Quercus

Q. floribunda = Q. dilatata
Q. formosana Greenidea nigra
Mollitrichosiphum fasciatum
Q. frainetto Phylloxera texana
(Hungarian or Italian Oak) Tuberculatus eggleri, etruscus
Q. frutex? Mexicallis analiliae ssp. pumilus
Q. gambelii (incl. hybrids, e.g. Lachnus allegheniensis
'undulata') Myzocallis discolor
(Gambel Oak) Neosymydobius chrysolepis
Stegophylla quercicola, quercifoliae
Thelaxes californica
Tuberculatus kiowanicus
Q. garryana Myzocallis punctata
(Oregon White Oak) Stegophylla essigi
Thelaxes californica
Tuberculatus annulatus,
columbiae, pallidus
Q. gemelliflora [Eutrichosiphum sp. (Noordam, in
prep.)]
Q. gilva Lachnus tropicalis
(Ichii-gashi) Metathoracaphis isensis
Quernaphis tuberculata
Q. glandulifera = Q. serrata
Q. glauca Allothoracaphis piyananensis
(Ara-kashi, Barin, Japanese Oak) Allotrichosiphum kashicola
Dermaphis japonensis
Eutrichosiphum dubium, [sp.
(Noordam, in prep.)]
Greenidea kuwanai, quercifoliae,
[sp. (Noordam, in prep.)]
Kurisakia querciphila
Lachnus tropicalis
Mollitrichosiphum glaucae,
nigrofasciatum
Neothoracaphis glaucae,
saramaoensis
Nipponaphis distyliicola, monzeni
Parathoracaphis setigera
Xenothoracaphis kashifoliae
Q. griffithi Eutrichosiphum khasyanum
Tuberculatus indicus, quercicola
Q. grisea Neosymydobius ajuscanus
Q. gunnisonii Lachnus allegheniensis
Myzocallis punctatus
Thelaxes californica
Tuberculatus kiowanicus
Q, havardii Phylloxera tuberculifera

Q. ilex Diphyllaphis mordvilkoi

(Holm Oak) Hoplocallis microsetosus,
microsiphon, pictus
Hoplochaetaphis zachvatkini
Lachnus iliciphilus, pallipes,
roboris
Myzocallis boerneri, castanicola,
cocciferina, mediterranea,
schreiben
Phylloxera ilicis, quercus
Siculaphis vittoriensis
Thelaxes dryophila, suberi,
valtadorosi
Tuberculatus annulatus,
pallidus
Q. ilicifolia Lachnochaitophorus querceus
(Bear Oak) Myzocallis bella, longiunguis,
spinosa
Q, imbricaria Myzocallis meridionalis, punctata,
(Shingle Oak) walshii
Q. incana Eutrichosiphum assamense,
(Ban Oak) garwhalense
Heminipponaphis querciphaga
Hoplocallis microsetosus
Lachnus acutihirsutus
Neothoracaphis garwhalensis,
yanonis
Q infectoria (incl. var. boissieri) Hoplocallis pictus
(Mountain Deciduous Oak) Hoplochaetaphis zachvatkini
Lachnus roboris
Myzocallis boerneri
Phylloxera quercus
Stegophylla sp.
Thelaxes suberi
Tuberculatus albosiphonatus,
maculipennis, moerickei,
pallescens, pallidus
396 Quercus

Q. ithaburensis Hoplocallis microsiphon, pictus

(Valonian Oak, Tabor Oak) Hoplochaetaphis zachvatkini
Lachnus crassicornis, swirskii
Myzocallis boerneri, glandulosa,
komareki, schreiberi
Phylloxera quercina
Thelaxes suberi
Q. kelloggii Phylloxera reticulata
(California Black Oak) Stegophylla essigi
Thelaxes californica
Tuberculatus annulatus, maureri,
pallidus
Q. laevis Myzocallis multisetis
(American Turkey Oak, Catesby
Oak)
Q. lanuginosa = Q. pubescens
Q. laurifolia Longistigma caryae
(Laurel Red Oak) Myzocallis granovskyi,
melanocera, pepperi, spinosa
[Phyllaphis nigra Ashmead, 1881]
Q. leucotriphora Serratocallis takahashii
Q. liaotungensis Lachnus siniquercus
Q. lobata Stegophylla essigi
(California White Oak, Valley Thelaxes californica
Oak) Tuberculatus annulatus,
californicus
Q. lusitanica Tuberculatus moerickei
Q. lyrata Myzocallis melanocera,
(Swamp White Oak) meridionalis
Neosymydobius memorialis
Q. macranthera Tuberculatus maximus
Q. macrocarpa Hoplochaitophorus quercicola
(Bur Oak, Mossycup Oak) Myzocallis bella, discolor,
punctata, rostropunctatus
Neosymydobius albasiphus,
canadensis, mimicus
Phylloxera querceti, similans
Stegophylla querci, quercicola
Thelaxes californica
Tuberculatus tuberculatus
Q. macrolepis Lachnus roboris
 Quercus 397

Myzocallis boerneri, komareki,

occidentalis, schreiberi
Thelaxes suberi
Q. marilandica Longistigma caryae
(Blackjack Oak) Myzocallis multisetis, punctata
Stegophylla quercicola
Q. mexicana Mexicallis ?calvus, spinifer
Neosymydobius ajuscanus
Q. michauxii Lachnochaitophorus obscurus
(Swamp Chestnut Oak) Longistigma caryae
Neosymydobius albasiphus
Q. microphylla Tuberculatus garciamartelli
Q. mongolica [Aphis gossypii]
(Mizu-nara, Mongolian Oak) Diphyllaphis konarae
Greenidea kuwanai
Lachnus tropicalis
[Macrosiphum kuricola
Matsumura, 1917]
Myzocallis castanicola
Neothoracaphis yanonis
Nipponaphis distyliicola
Phylloxera coccinea
Stomaphis japonica
Tuberculatus annulatus, borealis,
capitatus, higuchii, japonicus,
kashiwae, [naganoe (Shinji,
1941)], paiki, pappus,
paranaracola, querceus,
quercicola, querciformosanus,
stigmatus, yokoyamai
Q. morii Allothoracaphis piyananensis
Q. muehlenbergii Tuberculatus tuberculatus
(Chinquapin Oak)
Q. myrsinaefolia Allotrichosiphum kashicola
(Bamboo-leaved Oak, Shira-kashi) Greenidea kuwanai
Lachnus tropicalis
Neothoracaphis elongata,
querciphaga
Nipponaphis coreana
Parathoracaphis setigera
Q. nigra Lachnochaitophorus obscurus
(Water Oak, American Red Oak) Longistigma caryae
398 Quercus

Myzocallis granovskyi,
longiunguis, melanocera,
meridionalis, spinosa
Neosymydobius luteus
Q. nuttallii Myzocallis frisoni
Q. obtusiloba = Q. stellata
Q. pagoda = Q. falcata var. pagodifolia
Q. palustris Longistigma caryae
(Pin Oak, Swamp Spanish Oak) Myzocallis discolor, exultans,
frisoni, granovskyi, longiunguis,
melanocera, multisetis,
punctata, spinosa, walshii
Neosymydobius albasiphus
Patchia winforii
Q. paucidentata Neothoracaphis elongata
Q. peduncularis Neosymydobius butzei 'var.
papillata'
Tuberculatus mexicanus
Q. pedunculata = Q. robur
Q. persica Hoplochaetaphis zachvatkini
(Manna Oak) Lachnus swirskii ssp. persicae,
Lachnus sp.
Myzocallis komareki, persica
Thelaxes suberi
Tuberculatus maximus
Q. petraea Diphyllaphis mordvilkoi
(Bergek, Durmast Oak) Hoplochaetaphis zachvatkini
Lachnus pallipes, roboris
Moritziella corticalis
Myzocallis castanicola
Phylloxera coccinea, confusa,
foaae, glabra, italica, quercus
Stomaphis quercus
Thelaxes dryophila, suberi
Tuberculatus annulatus, eggleri,
borealis. neglectus
Q. phellos Myzocallis castanicola, exultans
(Willow Oak) Stegophylla quercicola
Tuberculatus querceus
Q. phylliraeoides Atarsaphis agrifoliae
(Ubame-gashi) Diphyllaphis alba
 Quercus 399

Eutrichosiphum tattakanum
Greenidea kuwanai, nipponica
Lachnus tropicalis
[Myzocallis punctata]
Tuberculatus pallidus, pilosus
Q. pontica Tuberculatus annulatus
Q. prinus Hoplochaitophorus heterotrichus,
(Chestnut Oak) quercicola
Myzocallis discolor, melanocera,
punctatus
Thelaxes californica
Q. pseudoturneri Tuberculatus borealis
Q. pubescens Diphyllaphis mordvilkoi
(Downy Oak) Hoplocallis microsiphon
Hoplochaelaphis zachvatkini
Lachnus iliciphila, roboris
Myzocallis castanicola,
mediterraneus, occidentalis
Phylloxera foaae, quercus
Thelaxes dryophila, suberi
Tuberculatus annulatus, eggleri,
inferus, maculipennis
Q. pyrenaica Diphyllaphis mordvilkoi
(Rebollo, Pyrenean Oak) Lachnus roboris
Myzocallis annulatus, castanicola,
eggleri, komareki, occidentalis,
schreiben
Tuberculatus remaudierei
Q. robur (incl. var. fastigiata) Acanthochermes quercus
(English Oak) Foaiella danesii
Hoplocallis microsiphon, ruperti
Hoplochaetaphis zachvatkini
Lachnus pallipes, roboris
Myzocallis bella, castanicola
Phylloxera coccinea, confusa,
foaae, glabra, italica, quercus
Stomaphis quercus
Thelaxes dryophila, suberi
Tuberculatus annulatus, borealis,
moerickei, neglectus, querceus
Q. rubra Diphyllaphis microtrema
(Northern Red Oak) Lachnochaitophorus querceus
Myzocallis bella, castanicola,
400 Quercus

exultans, granovskyi,
longirostris, longiunguis,
melanocera, multisetis, occulta,
punctata, spinosa, walshii
Stegophylla quercifoliae, quercina
Q. rugosa Mexicallis analiliae, areolatus,
(Netleaf Oak) spinifer
Tuberculatus mexicanus, spiculatus,
spiculatus ssp. rebecae
Q. salicifolia Myzocallis spinosa
Q. semicarpifolia Eutrichosiphum khasyanum, pyri
Myzocallis polychaeta
Q. serrata (incl. var. donarium) Cervaphis quercus
(Ko-nara) Diphyllaphis konarae, quercus
Eutrichosiphum davidi,
[narafoliae], pasaniae, taoi,
tattakanum
Greenidea kuwanai, quercifoliae
Kurisakia querciphila
Lachnus tropicalis
Mollitrichosiphum nigrofasciatum
Myzocallis castanicola, kuricola
Neothoracaphis garwhalensis,
quercicola, yanonis
Stomaphis japonicus
Tuberculatus capitatus, higuchii,
kashiwae, konaracola,
quercicola, stigmatus, yokoyamai
[ Tuberculatus fulviabdominalis,
kunugi, naganoe (Shinji, 1941)]
Q. sessiliflora = Q. petraea
Q. sessilifolia Neothoracaphis elongata
Q. shumardii Myzocallis elliotti, granovskyi,
(Shumard Red Oak) longiunguis, melanocera,
meridionalis, spinosa
Q. sinuata = Q. durandii
Q. spinosa Neothoracaphis tarakoensis
Q. stellata Hoplochaitophorus quercicola
(Iron Oak, Post Oak) Neosymydobius albasiphus,
quercihabitus
Phylloxera rileyi
Stegophylla quercifoliae
 Quercus 401

Q. stellata var. margaretta Myzocallis multisetis

(Sand Post Oak) Neosymydobius memorialis
?Phylloxera stellata
Stegophylla quercicola
Q. stenophylloides Eutrichosiphum tattakanum
(Taiwan-uraziro-gasi) Greenidea kuwanai
Metanipponaphis cuspidatae
Q. suber Hoplocallis pictus
(Cork Oak) Hoplochaetaphis zachvatkini
Lachnus pallipes
Myzocallis boerneri, [schreiberi]
Thelaxes suberi
Q. tomentella Tuberculatus pallidus
Q. trojana Apulicallis trojanae
(Macedonian Oak)
Q. turbinella Tuberculatus pallidus
Q. urbani var. parvifolia Myzocallis longirostris ssp.
tepehuaensis
Q. variabilis Cervaphis quercus
(Abe-maki, Gulcham, Korean Greenidea kuwanai, nipponica,
Oak) quercifoliae
Kurisakia querciphila
Lachnus tropicalis
Mollitrichosiphum nigrofasciatum
Myzocallis boerneri, castanicola,
kuricola
Neothoracaphis quercicola,
saramoensis
Phylloxera kunugi
Tuberculatus capilatus, kashiwae,
pallidus, quercicola,
querciformosanus, stigmatus
Q. velutina Hoplochaitophorus quercicola
(Black Oak, Quercitron Oak) Lachnochaitophorus querceus
Myzocallis bella, frisoni,
granovskyi, longiunguis,
melanocera, multisetis. walshii
Tuberculatus tuberculatus
Q. vicilurbe Hoplocallis microsiphon
Q. virginiana Lachnus allegheniensis
(Live Oak) Longistigma caryae
Moritziella sp. (Cuba)
402 Quercus

Myzocallis melanocera, spinosa

Phylloxera texana
Tuberculatus pallidus
Q. wislizeni Myzocallis agrifolicola
(Interior Live Oak) Stegophylla essigi, quercicola
Quercus spp. (unidentified) [Betacallis querciphaga]
Dermaphis crematogastri
Eutrichosiphum arunachali,
heterotrichum, izas, jugeshwari
manipurense, manoji,
narafoliae, neoalnicola,
nungsireiae, querciphaga,
raychaudhurii, russellae ssp.
lijiangshanense, simlaensis, taoi,
(apatii
Greenidea brachyunguis, haldari,
hangnigra, kumaoni, [mushana],
quercicola, querciphaga
[Lachnus chosoni Szelegiewicz,
1975]
[Longicaudus himalayensis HRL,
1965]
Longistigma xizangensis
Microunguis depressa
Mollilrichosiphum elongatum,
godavariense, luchuanum,
nigriabdominalis, niitakaensis,
shinjii, [tenuicorpus]
Myzocallis meridionalis, occultus
Neohormaphis calva
Neothoracaphis sutepensis
Nipponaphis querciphaga
Paranipponaphis takaoensis
Parathoracaphis elongata, gooti
Parathoracaphisella indica
[Phylloxera bipunctata]
Pseudothoracaphis himachali
[Reticulaphis distylii ssp. rotifera]
Schizoneuraphis malayna
[Stomaphis longirostris]
Thoracaphis arboris, flava
Tuberculatus cornutus, nervatus

Master key to the main groups of oak aphids

1 Body small, pear-shaped, broadest anteriorly. Antennae 3-segmented;
those of aptera with 1 rhinarium, those of alata with 2 rhinaria. Forewing
 Quercus 403

of alata without R s , and hindwing without oblique veins . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxeridae - KEY A (p. 403)
Body small or large, not broadest anteriorly. Antennae 3- to 6-segmented
but only 3-segmented in aleyrodiform apterae; always with 2 primary
rhinaria and, at least in alata, also with secondary rhinaria. Forewing of
alata with R s , and hindwing with 1-2 oblique veins . . . . . . . . . . . . . . 2
2 Aptera aleyrodiform, with head, thorax and first abdominal segment
fused (as prosoma), and reduced antennae, legs and posterior abdominal
segments. Alata with narrow annular secondary rhinaria, and forewing
with Cu1a and Cu lb touching or united at their bases . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hormaphidinae - KEY B (p. 406)
Aptera of normal aphid form with well-developed antennae and legs.
Alata with oval or transversely elongate secondary rhinaria and forewing
with Cu l a and Cu l b arising separately . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 SIPH tubular, much longer than basal width, with numerous hairs (Note:
SIPH tend to break off at or near bases in preserved specimens, so may
sometimes be missing in slide preparations) . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenideinae - KEY C (p. 411)
SIPH broadly conical, truncate, pore-like or absent, with or without
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Large aphids. First tarsal segments with 9 or more ventral hairs. SIPH
in form of large pores, often placed on broad conical bases clothed with
numerous hairs . . . . . . . . . . . . . . . . . . . . . . Lachninae - KEY D (p. 416)
Small to medium-sized aphids. First tarsal segments with 2-7 ventral hairs.
SIPH either truncate, small and pore-like, absent, or if on broad cones
then with at most a single ring of hairs . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 All adult viviparae alate . . . Drepanosiphinae (part) - KEY E (p. 418)
Colonies including apterae only or both apterae and alatae . . . . . . . . . .
. . . . . . . . . . . . . Drepanosiphinae (part) + Thelaxinae - KEY F (p. 430)

KEY A - Phylloxeridae on oaks

The taxonomy of the oak phylloxerids has been little worked, so the key
which follows is partial and tentative, and aims to summarize the available
knowledge rather than provide reliable determination to species. It refers
only to apterous females (not fundatrices after the first couplet), and is
based mainly on literature. Phylloxera texana, which has been shown to
host-alternate between Carya illinoensis and two Quercus spp., in Texas, could
not be included because the Quercus-feeding generations have not been
described.

1 In galls on leaves, with only two generations per year and no alatae;
fundatrices produce sexuales. Primary rhinarium on ANT III protruding
404 Quercus

almost to level of apex of PT, so that antenna appears to be double-tipped

(Fig. 90A). Immature fundatrix bearing stellate processes with 6-11
pointed secondary projections (Fig. 90B); processes of adult fundatrix
small, digitiform . . . . . . . . . . . . . . . . . . . . . . . . . Acanthochermes quercus
- Free-living on stems, leaves or roots. Primary rhinarium may protrude but
not nearly to level of apex of PT. Stellate processes if present also in adult
stage and with knob-like secondary projections . . . . . . . . . . . . . . . . . . . 2
2 Body with short, stellate, dorsal processes, bearing knob-like secondary
projections (Fig. 90C) . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera stellata*
- Dorsal processes if present simple, although often bearing bracts, spicules
or denticles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Dorsal processes either undeveloped or if present then very small, not or
hardly as long as their basal widths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Dorsal processes evident, mostly longer than their basal widths . . . . 6
4 Tergum with minute sclerotic ridges appearing as short, disconnected,
irregular lines. BL about 0.5 mm . . . . . . . . . . Phylloxera tuberculifera*
- Tergum nearly smooth. BL 0.70-0.85 mm . . . . . . . . . . . . . . . . . . . . . . . 5
5 Primary rhinarium on ANT III subapical (i.e. PT developed; Fig. 90D).
ABD TERG 2-5 with spiracles . . . . . . . . . . . . . . . . . . Phylloxera glabra
- Primary rhinarium on ANT I I I either absent or virtually apical (PT
undeveloped). ABD TERG 2-5 without evident spiracles . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera slanfordiana*
6 Dorsal processes constricted at extreme base, and bearing non-capitate,
usually pointed, apical hairs (Fig. 90E). (On roots) . . . Foaiella danesii
- Dorsal processes usually widened at extreme base, and their apical hairs
usually have expanded or capitate apices. (On aerial parts) . . . . . . . . 7
7 Longest dorsal processes on thorax clavate; i.e. constricted centrally and
with a swollen apex, or if short and knob-like or almost cylindrical they
are not much narrower at apex than at base (e.g. Figs 90F-H) . . . . . .
. . . Phylloxera quercus group (coccinea, foaae, ilicis, italicum, quercus)
- Longest dorsal processes on thorax tapering, without a constriction,
narrow distally; if cylindrical on distal part then much narrower near apex
than at base (e.g. Figs 90I-P) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Spicules on dorsal processes very small and arranged in spiral rows
(Figs 90I, J) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Spicules on dorsal processes larger and not arranged in spiral rows . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9 Longest dorsal processes more than 3 x their basal width, with a short
apical hair (Fig. 90I). ANT III long and thin. Dorsal cuticle with numer-
ous pits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera rileyi*
 Quercus 405

Fig. 90. A, ANT III of fundatrix of Acanthochermes quercus', B, stellate process of immature fundatrix of
A. quercus (after Grassi, 1912); C, stellate process of aptera of Phylloxera stellata (after Duncan,
1922); D, ANT III of aptera of Ph. glabra; dorsal processes of E, Foaiella danesii (after Grassi, 1912),
F-H, Phylloxera quercus group, I, Ph. rileyi (after Duncan, 1922), J, Ph. reticulata (after Duncan, 1922),
K, Ph. confusa (fundatrix, after Grassi, 1912), L, Ph. confusa (later generation aptera, after Grassi,
1912), M, Ph. kunugi (after Miyazaki and Teramoto, 1991), N, Ph. quercina, 0-P, Moritziella corticalis.

- Longest dorsal processes less than 3 x their basal width, with apical hair
long, 5-6 x its apical diameter (Fig. 90J). ANT III rather stout, broadest
on distal half. Dorsal cuticle finely reticulate . . . . Phylloxera reticulata*
10 Dorsal processes with broad, bract-like spicules (Figs90K, L). Apical
hairs on spinal processes long, at least 4 x their apical diameter ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera confusa*
- Dorsal processes with narrower, pointed or denticulate spicules, and with
short apical hairs which are usually less than 3 x their apical diameter
(except when the processes are very short) . . . . . . . . . . . . . . . . . . . . . . 11
11 All thoracic tergites and ABD TERG 1 with pleural as well as marginal
processes. Dorsal cuticle with nodulose ornamentation. Spiracles evident
on ABD TERG 2-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- Pleural processes absent from hind margin of pronotum and from ABD
TERG 1. Dorsal cuticle variously ornamented. Abdominal spiracles
present only on ABD TERG 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
406 Quercus

12 BL 2.0-2.5 mm. Dorsal processes conical, all less than 2 x their basal
diameters (Fig. 90M) . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxera kunugi
- BL 0.9-1.5 mm. Longest dorsal processes have attenuated, cylindrical
apices and are more than 3 x longer than their basal diameters (Fig. 90N)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxerina quercina
13 Rostrum short; when directed backwards not reaching to hind coxae.
Longest dorsal processes shorter than HT II. Dorsal cuticle with minute,
short ridges or wrinkles . . . . . . . . . . . . . . . . . . . . . . Phylloxera similans*
- Rostrum reaching well beyond hind coxae. Longest dorsal processes
usually longer than HT II; if a little shorter then the processes are pig-
mented and the dorsal cuticle is very rugose . . . . . . . . . . . . . . . . . . . . 14
14 Thoracic and anterior abdominal tergites with very rugose, denticulate
sculpturing. Dorsal processes pigmented, the longest usually not more
than 2 x HT II and often much shorter (Figs 90O, P) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Moritziella corticalis
- Dorsal cuticle less rugose, variously ornamented. Longest dorsal processes
often more than 2 x HT II, degree of pigmentation unknown . . . 15
15 Longest dorsal processes more than 0.8 x hind femora, which are rather
stout. Tibiae without annular ridges . . . . . . . . . . . Phylloxera querceti*
- Longest dorsal processes less than 0.7 x hind femora, which are relatively
slender. Tibiae with annular ridges . . . . . . . . . . . Phylloxera davidsoni*

KEY B - Hormaphidinae (Nipponaphidini) on oaks

The taxonomy of Nipponaphidini on oaks is poorly understood, despite the
revisionary work of A.K. Ghosh and Raychaudhuri (1973a, b). The uncertain-
ties will remain for most species until it is known whether they have primary
host forms, presumably forming galls on Distylium. Many nominal species
have been collected infrequently or only once, and host plants are often not
identified to species. Several species are recorded as feeding on other genera
as well as Quercus, especially Lauraceae (Litsea, Machilus). These records may
be of 'casual' occurrences, or due to errors of identification. Generic concepts
within this group are especially problematic.
The key to aleyrodiform apterae which follows is partial and simplified
to include only the better-known or more easily recognized species, but should
enable specimens to be placed in the right species group. The systematic part
of the book should be consulted for further information.

1 SIPH present as small pores or shallow cones on the posterior part of the
'abdominal plate' (= fused ABD TERG 2-7). Tarsi always 2-segmented
with claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- SIPH completely absent. Tarsi variably developed, reduced in some
species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
 Quercus 407

2 Prosoma completely fused with abdominal plate . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neohormaphis calva
- Prosoma at least showing partial separation from abdominal plate,
although the division may be indistinct medially (e.g. Fig. 91 A ) . . . . . 3
3 Dorsal cuticle of prosoma (head, thorax and first abdominal segment)
densely and entirely ornamented with nodules or pustules (Figs 91 A, B)
. . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Dorsal cuticle of prosoma with at least some areas devoid of pustules
(although the perpendicular marginal regions may be densely pustulate)
............................................................. 7
4 Prosoma ornamented with blunt conical nodules, often longer than their
basal widths (Fig. 91 A). Prosomal hairs very long and very fine . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis monzeni
- Prosoma ornamented with low pustules, all or mostly shorter than their
basal widths (e.g. Fig. 91B). Prosomal hairs short . . . . . . . . . . . . . . . . 5
5 Prosoma without tubercles. ABD TERG 7 with a pair of spinal hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nipponaphis distyliicola/coreana
- Prosoma with, in addition to the pustulate ornamentation, a series of
paired spinal and marginal tubercles. Posterior part of abdominal plate
(i.e. ABD TERG 7) without spinal hairs . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Spinal and marginal tubercles on prosoma thick and finger-like, with
broad bases. Prosomal hairs hardly longer than basal diameter of ANT
I I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Indonipponaphis tuberculata*
- Spinal and marginal tubercles very small. Prosomal hairs 2-3 x basal
diameter of ANT III . . . . . . . . . . . . . . . . . . . Indonipponaphis fulvicola*
7 Prosoma without any hairs. Antennae very short and unsegmented . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microunguis depressa*
- Prosoma with hairs. Antennae reduced and concealed beneath anterior
margin of prosoma between eyes, but 3-segmented . . . . . . . . . . . . . . . 8
8 Margins of prosoma densely pustulate. Prosomal hairs thick basally and
arising from prominent, somewhat tuberculate bases . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . Schizoneuraphis malayna/Nipponaphis querciphaga*
- Margins of prosoma pallisade-like. Prosomal hairs fine, without tuber-
culate bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 Abdominal plate with 5-10 spinopleural hairs each side of midline . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thoracaphis arboris
- Abdominal plate with only 1-2 spinopleural hairs on each side of midline
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xenothoracaphis kashifoliae
10 Prosoma completely fused with ABD TERG 2-7 . . . . . . . . . . . . . . . . 11
408 Quercus

- Prosoma at least showing partial (lateral) separation from ABD TERG

2-7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
11 Body flattened. Dorsal cuticle pale, not or only lightly sclerotized. Mar-
ginal hairs not spine-like. Dorsum with 4 pairs of long blunt or capitate
hairs, and 1 pair of similar hairs on ABD TERG 8 . . . . . . . . . . . . . . . .
............................................. 'Thoracaphis' flava
- Body box-like. Dorsal cuticle brown-black, sclerotized. Marginal hairs,
and 1-2 pairs on ABD TERG 8, spine-like . . . . . . . . . . . . . . . . . . . . . 12
12 Dorsal cuticle heavily sclerotized and corrugated, but not distinctly reticu-
late, with 5 pairs of spine-like spinopleural hairs, like the marginal hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parathoracaphis elongatus*
- Dorsal cuticle either granular with indistinct reticulation or heavily
sclerotized and strongly reticulate. Spinopleural hairs either long and fine,
minute or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Body almost circular. Dorsal cuticle granular with indistinct reticulation,
each prosomal segment bearing a pair of fine spinopleural hairs 40-
50 µm long. The 4 spine-like hairs on ABD TERG 8 in a single transverse
row, close together and all directed backwards (Fig. 91H) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parathoracaphis gooti
- Body narrow. Dorsal cuticle black, heavily sclerotized with strongly reti-
culate or convolute sculpturing, and with minute, inconspicuous spino-
pleural hairs. Spine-like hairs blunt, those on ABD TERG 8 not in a
transverse row, the more lateral pair not pointing backwards . . . . . 14
14 Dorsal cuticle strongly reticulate. ABD TERG 7 with 2 thick spine-like
marginal hairs of similar size close together on each side (Fig. 91F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parathoracaphis setigera
- Dorsal cuticle convoluted. ABD TERG 7 with 1 thick spine-like marginal
hair on each side, the spines anterior to it being smaller and apparently
on ABD TERG 6 (Fig. 91G) . . . . . . . . . . . Parathoracaphis kayashimai
15 Tarsi all 2-segmented, with claws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- Tarsi all rudimentary, 1-segmented and usually without claws, or absent
from fore and mid-legs and rudimentary on hind legs . . . . . . . . . . . 20
16 Entire prosoma, both dorsally and marginally, and also the abdominal
plate, ornamented with flat polygonal pustules (Fig. 91C). Prosomal hairs
all short and inconspicuous . . . . . . . . . . . . Paranipponaphis takaoensis
- Dorsal cuticle smooth or with different ornamentation. Prosomal hairs
long or short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 Prosoma only separate laterally from abdominal plate, which bears 6 pairs
of marginal hairs. Prosoma without evident sculpturing medially, except
around muscle plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
 Quercus 409

- Prosoma fully separated from abdominal plate, which is without marginal

hairs. Medial area of prosoma with reticulate, convolute or mosaic
sculpturing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18 Prosoma with a posterior pair of spinopleural hairs. Abdominal plate
with about 5 pairs of spinopleural hairs as well as the marginal ones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metathoracaphis isensis*
- Prosoma without a posterior pair of spinopleural hairs. Abdominal plate
with spinopleural hairs only on ABD TERG 7 (i.e. near posterior margin)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parathoracaphisella indica*
19 Prosoma and ABD TERG 8 with very long (100-150µm) marginal hairs
arising from tuberculate bases. ABD TERG 8 with 4 hairs. Entire dorsal
cuticle with fine, irregular, mosaic-like sculpturing (Fig. 91D) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allothoracaphis piyananensis
- Prosoma with marginal hairs absent or inconspicuous. ABD TERG 8 with
2 hairs. Dorsal cuticle with corrugated, convoluted sculpturing (Fig. 91E),
marginal areas of prosoma with reticulate sculpturing . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudothoracaphis himachali
20 Dorsal cuticle of prosoma and abdominal plate subdivided by clear, longi-
tudinal sutures into medial and lateral plates, which each carry charac-
teristic ornamentation. Margins of body with irregular lateral outgrowths
of venter (Fig. 91I) . . . . . . . . . . . . . . . . . . . . . . . . . . Atarsaphis agrifoliae
- Dorsal cuticle of prosoma and abdominal plate not subdivided . . . 21
21 Prosoma with long marginal hairs projecting around sides of body (which
in old adults is jet black and entirely opaque unless bleached before
mounting) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
- Prosoma with marginal hairs present or absent but not projecting around
sides of body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22 Marginal prosomal hairs flattened and expanded distally with toothed or
serrate apices (Fig. 48C) . . . . . . . . . . . . . . . . . Reticulaphis distylii group
- Marginal prosomal hairs with pointed apices (Fig. 91L) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermaphis spp.
23 Prosoma with rather thick, curved, acuminate spinopleural and sub-
marginal hairs. Prosoma and abdominal plate with pustulate ornamenta-
tion, and venter with smooth irregular lateral outgrowths (Fig. 91J). Hind
legs not projecting behind body . . . . . . . . . . . . . Quernaphis tuberculata
- Prosomal hairs absent or inconspicuous. Dorsal ornamentation not
pustulate. Hind legs usually projecting behind body . . . . . . . . . . . . . 24
24 Fore and mid-legs without tarsi, hind legs with rudimentary tarsi. Pleural
areas of prosoma with polygonal reticulate sculpturing, marginal areas
smooth (Fig. 91K) . . . . . . . . Neothoracaphis yanonis group (quercicola,
tarakoensis, yanonis)
 410 Quercus

Fig. 91. A, aptera of Nipponaphis monzeni with detail of central part of prosoma; detail of similar area
of prosoma of B, N. distyliicola, C, Paranipponaphis takaoensis, D, Allothoracaphis piyananensis,
E, Pseudothoracaphis himachali', dorsal (right) and ventral views of aptera with detail of spines on ABD
TERG 7 of F, Parathoracaphis setigera, 6, P. kayashimai; H, end of abdomen of aptera of
Parathoracaphis gooti; I, dorsal view of Atarsaphis agrifoliae; J, dorsal view of Quernaphis tuberculata',
K, dorsal (right) and ventral views of Neothoracaphis yanonis; L, marginal hairs of Dermaphis sp.
 Quercus 411

- All legs with rudimentary (1-segmented) tarsi. Cuticle of prosoma ridged,

corrugated or nearly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 Dorsal cuticle of prosoma with fine polygonal mosaic-like sculpturing,
apparently extending into marginal areas . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neothoracaphis garwhalensis*
- Dorsal cuticle of prosoma ridged, corrugated or nearly smooth . . . 26
26 Body elongate, BL 1.7-2.7 x maximum width. Prosoma smooth with a
distinct spinal ridge, which has a number of variably-developed constric-
tions and may have transverse furrows . . . . . . . . . . . . . . . . . . . . . . . . . 27
- Body oval, BL 1.4-1.5 x maximum width. Cuticle of prosoma corrugated
at least submarginally, spinal ridge absent or indistinct . . . . . . . . . . 28
27 Spinal ridge of prosoma with transverse furrows darkened and sclerotic,
separating 4 paler regions in between . . . . . . Neothoracaphis elongata*
- Spinal ridge of prosoma uniformly pale, with 4 prominent constrictions
but without dark furrows and pale intervening regions . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neothoracaphis saramoensis*
28 Prosoma of cleared specimens dark and distinctly corrugated over entire
dorsal surface . . . . . . . . . . . . . . . . . . . . . . . Neothoracaphis querciphaga*
- Prosoma of cleared specimens paler, and distinctly corrugated only sub-
marginally; mid-dorsal area darker, and smooth except for 3 transverse
furrows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 Hind legs pale . . . . . . . . . . . . . . . . . . . . . . . . . . . Neothoracaphis glaucae*
- Hind legs dark . . . . . . . . . . . . . . . . . . . . . . . . Neothoracaphis sutepensis*

Key C - Greenideinae on oaks (apterous viviparae unless otherwise stated)

Subtropical, oriental, evergreen oaks are the hosts of numerous species of
Greenideinae, but many of the aphid species are described from single collec-
tions, with the host unidentified to species. Some of the species separa-
tions in the key that follows are therefore rather tentative, particularly in
Eutrichosiphum and Mollitrichosiphum, and further work may reveal some
synonymies. The rather cumbersome subgeneric classifications that have been
erected for those two genera are probably best ignored for general purposes.

1 SIPH with short hairs. Body with very large, branched, hair-bearing pro-
cesses (Fig. 92A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cervaphis quercus
- SIPH with long hairs. Body without large branched hair-bearing processes
............................................................. 2
2 CAUDA usually with an apical papilla (Fig. 92B). SIPH always with pale
reticulation, at least at base often snapped off at base in preserved
specimens). Body often brown, very broadly pear-shaped and rather flat
(genus Greenidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- CAUDA never with an apical papilla. SIPH usually without any reticula-
tion. Body broadly pear-shaped, oval or elongate . . . . . . . . . . . . . . . . 8
412 Quercus

3 SIPH with pale reticulation extending over most of length (except at apex)
............................................................. 4
- SIPH reticulated only at base (subgenus Trichosiphum) . . . . . . . . . . . 5
4 ANT III reticulated over most of length except at extreme base, and also
distinctly imbricated distally . . . . . . . . . . . . . . . . Greenidea querciphaga
- ANT III neither reticulated nor imbricated . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea quercifoliae/(mushana)
5 SIPH very short and stout (Fig. 92D), less than 0.18 x BL and less than
0.75 x head width across (and including) eyes . . . . Greenidea haldari
- SIPH longer, more than 0.2 x BL, more than 0.8 x head width across
eyes (e.g. Fig. 92E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 R IV+V 2.4-3.0 x HT II . . . . . . . . . . . . . . . . . . . . Greenidea quercicola
- R IV+V 1.2-2.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Venter of abdomen strongly and densely spinulose. SIPH 0.29-0.36 x BL
(Fig. 92E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Greenidea kuwanai
- Venter of abdomen smooth or only locally or weakly spinulose, at least
on anterior sternites. SIPH 0.20-0.30 x BL . . . . Greenidea nipponica
8 Hind tibia with a series of narrow (stridulatory?) tranverse ridges spaced
out on basal 0.5-0.9 of length (e.g. Fig. 57D; genus Mollitrichosiphum)
............................................................. 9
- Hind tibia without transverse ridges (but often imbricated distally) . . . 15
9 Antenna 5- or 6-segmented. Dorsum pale. Hind tibia with over 50 trans-
verse, very narrow ridges extending about 0.9 of its length . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum niitakaensis
- Antenna 6-segmented. Dorsum pale or dark, or with a dark patch. Hind
tibia with only 14-30 transverse ridges on basal 0.5-0.6 . . . . . . . . . . 10
10 R IV+V 2.2-2.6 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- R IV+V 1.4-2.1 X HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Body pear-shaped, BL less than 2 x maximum width of abdomen . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum luchuanum
- Body elongate, BL more than 2 x maximum width of abdomen . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum godavariense
12 Body elongate, BL 2.3-2.5 x maximum width of abdomen. SIPH pale,
0.5-0.6 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Abdomen broader, BL about 2 x maximum width of abdomen or less.
SIPH dark, 0.3-0.4 x BL. (Dorsal abdomen with dark lateral areas linked
by a broad cross-band) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
 Quercus 413

Fig. 92. A, hair-bearing marginal process of aptera of Cervaphis quercus; B, CAUDA of Greenidea
quercifoliae; C, body hairs of aptera of Allotrichosiphum kashicola; SIPH of aptera of D, Greenidea
haldari, E, G. kuwanai, F, Eutrichosiphum tattakanum, G, Eu. heterotrichum (from type series); spinules
on dorsum of aptera of H, Eu. davidi and I, Eu. pasaniae; J, ANT III of aptera of Eu. dubium.
414 Quercus

13 Dorsal abdomen with a central dark patch . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum nigriabdominalis*
- Dorsal abdomen without a central dark patch . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum shinjii
14 R IV+V 1.5-1.8 x HT II . . . . . . . . Mollitrichosiphum nigrofasciatum
- R IV+V 1.4-1.6 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . Mollitrichosiphum nigrofasciatum ssp. glaucae
15 Antenna 5-segmented, rarely 6-segmented in some specimens . . . . . 16
- Antenna 6-segmented, rarely 5-segmented . . . . . . . . . . . . . . . . . . . . . . . 31
16 Longest hairs on antennae and dorsum distinctly capitate (Fig. 92C) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Hairs on antennae and dorsum with pointed, blunt, expanded or bifurcate
apices, but never capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 SIPH about 0.28 x BL. R IV+V about 2 x HT II . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allotrichosiphum assamense
- SIPH more than 0.5 x BL. R IV+V about 1.7 x HT II . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allotrichosiphum kashicola
18 Abdominal tergum with numerous spinules scattered or in irregular rows
over entire surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Abdominal tergum at least with mesial area devoid of spinules . . . 23
19 Abdominal tergum with densely scattered spinules which are much longer
than their basal diameters (Fig. 92H). SIPH only about 0.2 x BL or less.
R IV+V 2.9-3.1 x HT II . . . . . . . . . . . . . . . . . . Eutrichosiphum davidi
- Abdominal tergum with small spinules, hardly longer than their basal
diameters, in numerous irregular rows (e.g. Fig. 92I). SIPH 0.23-0.42 x
BL. R IV+V 1.5-2.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Dorsum pale. R IV+V 1.5-1.7 x HT II. SIPH pale at base, black distally,
0.40-0.55 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum shiicola
- Dorsum pale or dark. R IV+V 1.8-2.5 X HT II. SIPH uniformly dark
and 0.22-0.28 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Dorsal abdominal hairs all with acute apices . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum arunachali*
- Dorsal abdominal hairs mostly with blunt or furcate apices . . . . . . 22
22 Dorsum brown-black, R IV+V 1.8-2.3 x HT II . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pasaniae
- Dorsum mainly pale, sometimes dark at margins. R IV+V 2.2-2.9 x HT
II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pseudopasaniae
 Quercus 415

23 SIPH more than 0.4 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

- SIPH less than 0.4 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
24 R IV+V 3.0-3.6 x HT II . . . . . . . . . . . . . Eutrichosiphum quercifoliae
- R IV+V 1.3-1.7 X HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 ANT PT/BASE ,1.4-1.7. Abdominal tergum entirely devoid of spinules,
with a brown central patch on ABD TERG 2-4. Dorsal abdominal hairs
with blunt apices . . . . . . . . . . . . . . . . . . . . Eutrichosiphum manipurense*
- ANT PT/BASE 1.0-1.2. Abdominal tergum entirely pale, with only
mesial area devoid of spinules. Dorsal hairs mostly with furcate apices
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum jugeshwari *
26 R IV+V 2.9-3.5 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
- R IV+V 1.1-2.6 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
27 SIPH pale with dusky apices, and 3.8-4.3 x their maximum width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum assamense
- SIPH brown-black, barrel-shaped, 2.5-2.8 x their maximum width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum nungsireiae *
28 R IV+V only 1.17-1.35 x HT II. Dorsal abdominal hairs mostly with
blunt or acute apices, never furcate . . . . . . . . . . Eutrichosiphum tapatii
- R IV+V 1.9-2.6 x HT II. Dorsal abdominal hairs with blunt, acute or
furcate apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 Body elongate, with BL more than 2 x maximum width. SIPH pale
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphon taoi *
- Body pear-shaped, BL less than 2 x maximum width. SIPH pale or dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 Most dorsal abdominal hairs with distinctly bifurcate or multifurcate
apices . . . . . . . . . . . . . . . . . . . . Eutrichosiphum garwhalense/simlaensis*
- Dorsal abdominal hairs mostly with blunt, acute or slightly expanded
apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum pyri *
31 SIPH with hairs mostly long, those on middle part being all of similar
length and thickness (e.g. Fig. 92F); some basal and subapical hairs may
be shorter, but these are not interspersed with much longer hairs . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
- SIPH with hairs of very different lengths interspersed throughout (varying
by a factor of 2 or more; Fig. 92G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
32 Hairs on SIPH all with acute apices, occasionally 1-2 bifurcate (e.g.
Fig.92F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
- Several of hairs on basal part of SIPH with furcate apices . . . . . . . 34
33 Dorsal abdomen with a brown-black quadrate patch centred on ABD
TERG 3-5. SIPH 0.32-0.36 x BL . . . . Eutrichosiphum querciphaga*
416 Quercus

- Dorsal abdomen usually dark laterally with a pale anteriomesial area.

SIPH 0.17-0.32 x BL . . . . . . . . . . . . . . . . Eutrichosiphum tattakanum
34 R IV+V 1.6-2.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum garwhalense*/raychaudhurii*
- R IV+V 2.3-3.1 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
35 SIPH 0.17-0.20 x BL . . . . . . . . . . . . . . . . . . . . Eutrichosiphum flavum
- SIPH 0.22-0.29 x BL . . . . . . . . . . . . . . . . . Eutrichosiphum khasyanum
36 SIPH 3.7-7.5 x their maximum width, and 0.23-0.40 x BL . . . . . 37
- SIPH 2.8-3.0 x their maximum width, and at most 0.21 x BL . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
37 ANT III smooth . . . . . . . . . . . . . . . . . . . . Eutrichosiphum heterotrichum
- ANT III imbricated (Fig. 92J) . . . . . . . . . . . . . . Eutrichosiphum dubium
38 ANT III with 17-19 hairs, mostly more than 2 x basal diameter of seg-
ment. ABD TERG 7 with 15-18 hairs . . . . . Eutrichosiphum russellae
- ANT III with 9-10 hairs, of which only 2-3 are more than 2 x basal
diameter of segment. ABD TERG 7 with 10 or fewer hairs . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eutrichosiphum rameshi*

KEY D - Lachninae on oaks (apterous viviparae except where otherwise

stated)
1 Rostrum much longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Rostrum shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 ABD TERG 1-7 all with paired dark spinal patches, sometimes broken
into clusters of small sclerites . . . . . . . . . . . . . . . . . . . Stomaphis quercus
- ABD TERG 3-5 never with paired dark spinal patches, and ABD TERG
1, 2 and/or 6 either without patches or with small clusters of sclerites
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis japonica
3 BL more than 5 mm. Forewings of alata not maculate, and with an
elongate pterostigma that curves around tip of wing . . . . . . . . . . . . . . 4
- BL almost always less than 5 mm. Forewings of alata (where known)
maculate, with a normal, blunt pterostigma . . . . . . . . . . . . . . . . . . . . . . 5
4 HT II 1.8-2.3 x HT I . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- HT II about 2.6 x HT I . . . . . . . . . . . . . . . . Longistigma xizangensis*
5 SIPH pores on very flat cones, hardly raised above level of dorsum, with
a broad, reticulated flange, and little or no pigmentation (Fig. 93A).
Appendages and entire body, dorsally as well as ventrally, clothed in a
dense pile of hairs; those on ABD TERG 1-5 with bases less than 20 µm
apart . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus crassicornis
 Quercus 417

- SIPH pores on conical bases that are usually large, pigmented and clearly
demarcated, but sometimes small and pale; flange small. Hairs numerous,
but not forming a dense pile; hair bases more than 20 µm apart, at least
on dorsum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 R IV+V 1.20-1.31 x HT II . . . . . . . . . . . . . . . . . . . Lachnus takahashii
- R IV+V 0.65-1.12 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 R IV+V 0.65-0.82 x HT II, but if 0.79-0.82 x HT II then less than 0.2
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- R IV+V 0.80-1.12 x HT II; if less than 0.85 x HT II then more than
0.2 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8 Maximal basal diameter of SIPH cone 0.16-0.24 mm. ANT PT/BASE a
little less than 0.5. R IV+V 0.74-0.82 x HT II. Hairs on ABD TERG
1-3 all shorter than basal diameter of ANT III . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus acutihirsutus
- Maximal basal diameter of SIPH cone 0.3-0.7 mm. ANT/PT BASE 0.5
or more. R IV+V 0.65-0.70 x HT I I . Longest hairs on ABD TERG 1-3
more than 1.5 x basal diameter of ANT III . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus allegheniensis
9 Hind tibia with some of more distal hairs on ventral side very long and
fine-pointed, at least 2-3 x longer than neighbouring short hairs
(Fig. 93D). Mesosternal processes absent or only weakly developed. SIPH
cones small and often pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Hind tibia with hairs on ventral side often longer and finer distally, but
not with some hairs 2-3 times longer than others. Mesosternum of summer
aptera with a pair of mammariform or papilliform processes. SIPH cones
usually large and pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 Dorsal body hairs long and fine-pointed . . . . . . . . . . . Lachnus pallipes
- Dorsal body hairs very short (less than 30 µm) and blunt . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . Lachnus sp. ex Quercus aegilops, Turkey
11 SIPH cones small and pale. Hairs on ABD TERG 1-6 all short and blunt,
maximally about 20 µm . . . . . Lachnus sp. ex Quercus persica, Turkey
- SIPH cones usually large (basal diameter more than 0.3 mm) and well-
pigmented. Hairs on ABD TERG 1-6 variable, but the longest always
more than 20 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 ANT PT/BASE 0.5 or more. Mesosternal processes mammariform
(Fig. 93B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus tropicalis
- ANT PT/BASE clearly less than 0.5. Mesosternal processes papilliform
(Fig. 93C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 ANT PT shorter than diameter of primary rhinarium on VI (Fig. 93E).
Hairs on ABD TERG 1-3 mostly thick, blunt and less than 30 µm long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus swirskii
418 Quercus

Fig. 93. K, SIPH of aptera of Lachnus crassicornis, dorsal and side views; mesosternal processes of
apterae of B, L. tropicalis and C, L. roboris; D, distal part of hind tibia of L. pallipes; ANT VI of
E, L. swirskii and F, L roboris.

- ANT PT longer than diameter of primary rhinarium on VI (Fig. 93F).

Hairs on ABD TERG 1-3 variable but mostly pointed, and mostly more
than 30 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus roboris

KEY E - Drepanosiphinae on oaks (alate viviparae only)

Aphids of the subfamily Drepanosiphinae which exist only as alatae in their
parthenogenetic phase comprise mainly two large genera, Tuberculatus
and Myzocallis. A few Myzocallis have apterous as well as alate viviparae
and are therefore in Key F. Within both these genera there are some distinct
subgeneric groupings, which are referred to in the key. Hoplocallis is
sometimes treated as a subgenus of Myzocallis, but is here given separate
generic status.
 Quercus 419

1 Dorsal abdomen with one or more spinal processes (genus Tuberculatus)

............................................................. 2
- Dorsal abdomen without spinal processes . . . . . . . . . . . . . . . . . . . . . . . 44
2 Dorsal abdomen with one large black bifurcate spinal process on ABD
TERG 3 only (Fig. 94A) . . . . . . . . . . . . . . . . . . . . Tuberculatus querceus
- Dorsal abdomen with more than one pair of spinal hair-bearing processes,
separated or only united basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Each tibia with a conspicuous dark brown to black spot at its base
(subgenus Pacificallis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Tibiae pale or variably pigmented but never with brown-black basal spots
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
4 Forewing with Cu1b thickly bordered with fuscous, the width of the vein
plus borders at its half-way point being 0.05-0.07 mm . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus maureri
- Forewing with the Cu1b vein itself often thicker and darker than other
veins, but not thickly bordered; vein plus borders not more than 0.035 mm
wide at its half-way point . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Spinal hairs on pronotum less than 10 µm long. Spinal processes absent
from thorax or at most developed as tubercular hair-bases shorter than
their basal diameters . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus passalus
- Spinal hairs on pronotum more than 10 µm long. Spinal processes devel-
oped on pronotum and mesonotum, with at least those on mesonotum as
long as their basal diameters and often finger-like . . . . . . . . . . . . . . . . 6
6 Pronotum with only 1 evident (posterior) pair of spinal processes
(although the anterior prothoracic spinal hairs may have flat tuberculate
bases) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Pronotum with 2 pairs of evident spinal processes, both pairs longer than
their basal diameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7 Head with 6-13 dorsal hairs between the eyes. Pronotum with anterior
marginal hairs. Embryos with marginal hairs on ABD TERG 1-4
duplicated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus californicus
- Head with 4 dorsal hairs between eyes. Prothorax without anterior
marginal hairs. Embryos with marginal hairs on ABD TERG 1-4 single
............................................................. 8
8 ANT BASE VI with 1 hair. Embryos and immatures with most dorsal
hairs strongly capitate, usually not on sclerites . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus columbiae
- ANT BASE VI with 2 hairs. Embryos and immatures with most dorsal
hairs rod- or spear-shaped with blunt or pointed apices, those of immatures
usually arising from pigmented sclerites . . . . Tuberclatus chrysolepidis
420 Quercus

9 ANT III with 7-14 secondary rhinaria. ABD TERG 1-4 with spinal pro-
cesses all of similar size . . . . . . . . . . . . . . . . . . . Tuberculatus kiowanicus
- ANT III with 1-7 secondary rhinaria. ABD TERG 1 and 2 with spinal
processes about twice as tall as those on more posterior tergites . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 R IV+V 0.11-0.13 mm with usually 2 pairs of lateral accessory hairs as
well as some ventral hairs. Head with anteriodorsal hairs slightly but
distinctly capitate . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus quercifolii
- R IV+V 0.095-0.11 mm, usually with 1 pair of lateral accessory hairs as
well as some ventral ones. Head with anteriodorsal hairs blunt but not
distinctly capitate . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus pallidus
11 First tarsal segments all with 5 ventral hairs (as well as 2 dorsal ones).
Immatures with spiculose dorsal hairs (Fig. 94B; spicules also discernible
on the longest dorsal hairs of embryos within adult alata) (subgenus
Toltecallis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- First tarsal segments all or mostly with 6 (or 7) ventral hairs. Dorsal hairs
of immatures and embryos without spicules . . . . . . . . . . . . . . . . . . . . . 14
12 ANT PT/BASE more than 1.5, and PT more than 1.6 x R IV+V. ANT
II pigmented. (ABD TERG 1-4 with spinal processes) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus mexicanus
- ANT PT/BASE less than 1.3, and PT less than 1.4 x R IV+V. ANT II
pale. (ABD TERG 1-2 or 1-4 with spinal processes) . . . . . . . . . . . . . 13
13 R IV+V 0.092-0.101 mm. ABD TERG 1-2 (and also mesonotum) with
spinal processes; height of those on mesonotum 0.3-0.5 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus garciamartelli
- R IV+V 0.112-0.162 mm. Mesonotum, ABD TERG 1-4, and often 6 also,
with spinal processes; those on mesonotum 0.7-1.1 x HT II . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus spiculatus*
14 Frontal hairs mostly shorter than, and hairs on ANT III all much shorter
than, basal diameter of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Frontal hairs usually 2 or more x basal diameter of ANT III, and longest
hairs on ANT III often as long as basal diameter or longer . . . . . . 26
15 Forewings with a pattern of infuscation (use hand lens). ABD TERG 1
with a pair of large pigmented spinal processes, those on more posterior
tergites being much smaller . . . . . . . . . . . . . . Tuberculatus tuberculatus
- Forewings mainly pale. Spinal processes on ABD TERG 1 often unpig-
mented, and never larger than those on ABD TERG 2-3 (subgenus Tuber-
culoides) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 ABD TERG 1-4 each with a pair of pale spinal processes (e.g. Fig. 94C;
although those on 4 may be small) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
 Quercus 421

Fig. 94. A, spinal process of Tuberculatus querceus; B, spiculose hairs of immature T. mexicanus;
spinal processes of C, T. borealis and D, T. neglectus (after Krzywiec, 1965).

- Only ABD TERG 1-3 with spinal processes (e.g. Fig. 94D) . . . . . . . 20
17 R IV+V 0.13-0.19 mm, 1.15-1.50 x HT II, with 3 pairs of lateral
accessory hairs (as well as 3-6 ventral ones) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus remaudierei
- R IV+V 0.08-0.12 mm, 0.80-1.05 x HT II, with 2 pairs of lateral acces-
sory hairs (and 3-5 ventral ones) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Spinal hairs of embryos (seen through abdominal cuticle of alate vivipara)
all similar in length to the long marginal hairs on the same tergites
(Fig. 95A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus eggleri
- Spinal hairs on at least ABD TERG 2-3 of embryos only half as long as
marginal hairs on same segments or shorter . . . . . . . . . . . . . . . . . . . . 19
19 Embryos with spinal hairs on ABD TERG 1-6 all about 5-8 µm long, thin,
with blunt apices; marginal hairs 15-45 µm long, with those on at least
ABD TERG 2-3 not overlapping those of next tergite (Fig. 95B) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus borealis
- Embryos with distinctly capitate spinal hairs, those on ABD TERG 1
being 6-10 µm long, those on more posterior tergites 26-75 µm; marginal
hairs 26-75 µm long, all except those on ABD TERG 1 overlapping those
on next tergite (Fig. 95C) . . . . . . . . . . . . . . . . . . . Tuberculatus africanus
20 Spinal hairs of embryos mostly very long and overlapping between
segments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
- Spinal hairs on ABD TERG 1-5 of embryos all very short (less than 5 µm)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
21 ANT PT/BASE 2.1-2.8. R IV+V 1.20-1.35 X HT II and bearing 12-15
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus etruscus
- ANT PT/BASE 0.9-1.8. R IV+V 0.90-1.15 X HT II and bearing 7-10
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
422 Quercus

Fig. 95. Dorsal chaetotaxy of embryos of A, Tuberculatus eggleri, B, T. borealis, C, T. africanus,

D, T. annulatus (after Richards, 1969a), E, T. neglectus.

22 BL 1.3-2.3. ANT PT/BASE 1.42-1.78 . . . . . . . . . Tuberculatus inferus

- BL 2.0-3.1. ANT PT/BASE 0.88-1.39 . . . . . . . Tuberculatus maximus
23 SIPH almost completely pale. ANT PT/BASE 0.85-0.95. ANT III usually
with dusky section about one third from base (use hand lens) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus albosiphonatus
- SIPH dark on apical third or more. ANT PT/BASE 0.87-1.53, but if less
than 0.95 then ANT III is pale except for black distal section . . . . 24
24 ANT PT/BASE 0.87-1.14. ANT III pale except for black distal section.
ABD TERG 2-5 of embryos with marginal hairs 8-13 µm long, like the
spinal hairs on same segments; usually only ABD TERG 8 with long
capitate hairs (Fig. 95D) . . . . . . . . . . . . . . . . . . . . Tuberculatus annulatus
- ANT PT/BASE 0.95-1.53, but if less than 1.14 then ANT III usually has
a dusky section around the most distal secondary rhinarium. Embryos
with marginal hairs on ABD TERG 2-5 always much longer than spinal
hairs on those segments, and at least ABD TERG 7 as well as 8 with long
capitate hairs (Fig. 95E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 ANT PT/BASE 0.95-1.13. ANT III usually with a dusky central section.
SIPH a little shorter than R IV+V . . . . . . . . . . Tuberculatus moerickei
- ANT PT/BASE 1.14-1.53. ANT III pale except for black distal section.
SIPH a little longer than R IV+V . . . . . . . . . . . Tuberculatus neglectus
26 Thorax of mounted specimens pale (pronotum sometimes marginally
dark). Secondary rhinaria without ciliate rims. Forewing veins not ending
in blackish triangles (subgenus Orientuberculoides) . . . . . . . . . . . . . . 27
- At least mesothorax usually dark brown dorsally and/or ventrally or with
a dark brown dorsal patch, but if pale then secondary rhinaria have ciliate
rims and all wing veins end in blackish triangles . . . . . . . . . . . . . . . . 35
 Quercus 423

27 Spinal processes on ABD TERG 1-3 low, conical or rounded, hardly

longer than their basal diameters. Pronotum without spinal processes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus kashiwae
- Spinal processes at least on ABD TERG 3 usually finger-like, clearly
longer than their basal diameters. Pronotum usually with at least one pair
of spinal processes; if with none, then processes on ABD TERG 1-2 are
2.5-3.5 x their average width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 Pronotum only with a single (posterior) pair of (sometimes very small)
spinal processes. Hairs on ANT III not deeply pigmented . . . . . . . . 29
- Pronotum with 2 pairs of spinal processes. Longest 1-3 hairs on ANT III
capitate and often markedly pigmented, the sockets around the hair-bases
also being pigmented. Similar hairs often present on ANT I and II and
on frons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
29 Head with 3 pairs of long capitate dorsal hairs anterior to the lateral ocelli
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus querciformosanus
- Frons with only 2 pairs of long capitate dorsal hairs anterior to the lateral
ocelli, plus 1 much shorter pair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 Spinal processes on ABD TERG 3 wholly pigmented and often with pig-
ment extending between their bases, contrasting with pale processes on
ABD TERG 1 and 2. SIPH with distal half black . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus konaracola
- Spinal processes on ABD TERG 3 pale or only dusky on distal half. SIPH
pale or dark only on distal third or less . . . . . . . . . . . . . . . . . . . . . . . . 31
31 Embryos with all spinal hairs on ABD TERG 2-6 short, up to half of
length of marginal hairs on same segments . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus paranacola
- Embryos with spinal hairs on ABD TERG 2-6 long, comparable in length
to marginal hairs on same segments . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
32 Hairs on anterior side of ANT III (opposite rhinaria) short, thin and
cylindrical or club-shaped, only up to about 0.6 x basal diameter of
segment, and not on raised bases. (ANT III with 3-6 secondary rhinaria,
confined to basal 0.3-0.5 of segment even in spring populations. Fore-
wing with Cu la , Cu l b and media all of similar thickness and degree of
pigmentation) .............................. Tuberculatus higuchii
- Hairs on anterior side of ANT III capitate, 0.75-1.50 x basal diameter
of segment, with raised bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
33 Longest hairs on ANT III 0.75-1.20 x basal diameter of segment. ANT
III in spring populations with 4-10 secondary rhinaria extending over
0.5-0.9 of segment, only in midsummer specimens confined to basal 0.3
of segment and 2-4 in number. Cu l a and Cu lb of forewing thicker and
darker than distal branches of media (use hand lens) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus yokoyamai
424 Quercus

- Longest hairs on ANT III 1.3-1.5 x basal diameter of segment. ANT III
with 4-7 secondary rhinaria restricted to basal 0.5 of segment even in
spring populations. Cu la , Cu lb and media of forewing all of similar
thickness and degree of pigmentation . . . . . . . . . . . . Tuberculatus paiki
34 Front of head, ANT I-III and dorsal thorax all with strongly capitate
hairs. Dorsal hairs on basal half of hind tibia rather stiff, about 1.0-1.3
x diameter of tibia, with blunt or slightly capitate apices. Embryos with
spinal hairs on ABD TERG 2-6 very thickly capitate . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus capitatus
- Front of head, ANT I-III and dorsal thorax with acute, blunt and/or
slightly capitate hairs. Dorsal hairs on basal half of hind tibia thin with
very fine apices, up to 2 x diameter of tibia. Embryos with spinal hairs
on ABD TERG 2-6 blunt or only slightly capitate . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus fangi
35 Longest hair on ANT II 2 or more x longer than longest hair on ANT
III. Pronotum without any spinal processes (subgenus Camelaphis) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
- Longest hair on ANT II usually similar in length to or shorter than longest
hair on ANT III, or not more than 1.5 x longer. Pronotum with or
without spinal processes (subgenus Acanthocallis) . . . . . . . . . . . . . . . 38
36 Mesonotum pale. Abdomen without dark marginal processes. ABD
TERG 1-3 with mainly pale, sometimes black-tipped, spinal processes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus pallescens
- Mesonotum dark. Abdomen with large, dark, spiculose marginal pro-
cesses, those on ABD TERG 3 being about as long as the siphunculi.
Spinal processes also dark, at least on ABD TERG 2 and 3 . . . . . . 37
37 Forewings maculate. Spinal processes on ABD TERG 1-4 pigmented, and
shorter, pale spinal processes also present on more posterior tergites.
SIPH only dark basally . . . . . . . . . . . . . . . . . Tuberculatus maculipennis
- Forewings not maculate. Abdomen with 2 pairs of dark spinal processes
(very long and basally joined on ABD TERG 3). SIPH wholly dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus cornutus
38 Forewings with many fine hairs, not only on posterior margin of ptero-
stigma but also scattered over most of wing membrane distal to Cu lb
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
- Forewings without hairs on wing membrane . . . . . . . . . . . . . . . . . . . . 40
39 All forewing veins evenly and broadly bordered with fuscous over their
entire length. Long hairs on inner side of ANT III very finely pointed.
Spinal processes on ABD TERG 1-3, especially those on 3, much larger
and darker than those on ABD TERG 4-5 . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus quercicola
 Quercus 425

- Forewings with Cu la and Cu l b and unbranched base of media fuscous

bordered, but distal branches of media unbordered. Hairs on inner side
of ANT III with acute or capitate apices. Spinal processes on ABD TERG
1-3 short; those on 3 not much longer than their basal diameters
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus grisipunctatus*
40 Pronotum with 2 pairs of (pale) spinal processes . . . . . . . . . . . . . . . . 41
- Pronotum without spinal processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
41 Mesonotum without spinal processes . . . . . . . . Tuberculatus nervatus*
- Mesonotum with a pair of spinal processes . . . . . . . . . . . . . . . . . . . . . 42
42 Frontal hairs strongly capitate. Dorsal hairs on femora and on basal half
of hind tibia all stiff and distinctly capitate (except ssp. radisectuae) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus japonicus
- Frontal hairs with acute or rounded apices, sometimes slightly capitate.
Dorsal hairs on femora pointed, and many of those on basal half of hind
tibia fine and pointed (a few may be blunt or slightly capitate) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus indicus
43 Forewing veins with broad fuscous borders. Hind tibia mainly pale. ANT
III with 12-24 secondary rhinaria . . . . . . . . . . . . . Tuberculatus pilosus
- Forewing veins not bordered. Hind legs entirely black, in sharp contrast
to other legs. ANT III with 3-8 secondary rhinaria . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tuberculatus stigmatus
44 Head and prothorax with a median longitudinal pale stripe extending
backward from the median ocellus between bands of pigment (Fig. 96A).
Pronotum with anterior and posterior spinal and lateral clusters of small
hairs. Abdomen with paired dark spinal sclerites that are fused posteriorly
across midline, on ABD TERG 1-7 (Fig. 96A), or on 1-2 only. ANT
PT/BASE 0.50-0.95 (genus Hoplocallis) . . . . . . . . . . . . . . . . . . . . . . . . 45
- Head and prothorax pale or variously pigmented but never with a pale
median longitudinal stripe. Pronotal hairs not in clusters, and often absent
anteriolaterally. Abdomen either without sclerites or with paired sclerites
not usually fused across the midline, at least on anterior tergites. ANT
PT/BASE 0.95-3.50 (genus Myzocallis) . . . . . . . . . . . . . . . . . . . . . . . . 48
45 R IV+V stilletto-shaped, more than 1.75 x HT II. Abdomen with dark
sclerites only on ABD TERG 1 and 2, and with segmental clusters of
pleural hairs on ABD TERG 1-5 . . . . . . . . . . Hoplocallis microsetosus
- R IV+V less than 1.65 x HT II. Abdomen usually with dark sclerites on
ABD TERG 1-7 (fused across midline at least on 1-5; Fig. 96A). ABD
TERG 1-5 without clusters of pleural hairs . . . . . . . . . . . . . . . . . . . . . 46
46 SIPH small, c. 0.04 mm, 0.2-0.3 x R IV+V and about 0.5 x as long
as knob of CAUDA. R IV+V stilletto-shaped, 1.30-1.65 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplocallis microsiphon
426 Quercus

Fig. 96. A, dorsal body markings of Hoplocallis pieta; B, ANT III of Myzocallis kuricola; C, CAUDA and
anal plate of M. granovskyi; D, body and wing markings of M. bella; E, head and prothorax of
M. melanocera; spinal sclerites on ABD TERG 4 of F, M. komareki and G, M. cocciferina; pterostigma
of H, M. schreiberi and I, M. komareki.

- SIPH 0.07-0.10 mm, 0.50-0,85 x R IV+V, about as long as knob of

CAUDA. R IV+V obtuse, 1.00-1.55 x HT II . . . . . . . . . . . . . . . . . . 47
47 ANT PT/BASE 0.5-0.8. R IV+V 1.25-1.55 x HT II. ANT III with 2-4
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplocallis pictus
- ANT PT/BASE 0.80-0.95. R IV+V 1.0-1.2 x HT II. ANT III with 3-5
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplocallis ruperti
 Quercus 427

48 Forewings with pigmentation thickly bordering all veins, or extending

between veins (sometimes not so evident in spring populations) . . . 49
- Forewings without pigmentation along or between veins (except along
costal margin, and sometimes Rs and/or Cu1b) . . . . . . . . . . . . . . . . . . 53
49 ANT III with long pointed anteriorly-directed hairs, 2 or more x basal
diameter of segment (Fig. 96B). ANT PT/BASE about 1. Forewing
pigmentation faint and weakly delimited, strictly bordering veins . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis kuricola
- ANT III with all hairs shorter than basal diameter of segment. ANT
PT/BASE 1.0-3.5. Forewing pigmentation usually heavy, extending
between veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
50 SIPH dark, on large dark sclerites, with or without spicules . . . . . . 51
- SIPH pale, with spiculose ornamentation . . . . . . . . . . . . . . . . . . . . . . . 52
51 SIPH (and basal sclerites) with rows of spicules. Forewing pigmentation
patchy, discontinuous, between veins . . . . . . . . . . . . Myzocallis discolor
- SIPH (and basal sclerite) smooth. Forewing pigmentation heavy, mainly
following veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis agrifolicola
52 R IV+V less than 0.11 mm long, less than 0.75 x ANT VI BASE . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis punctata
- R IV+V more than 0.11 mm long, more than 0.8 x ANT VI BASE
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis mimica*
53 Head and pronotum with a distinct dark, median longitudinal stripe
(Fig. 35E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis castanicola
- Head and pronotum variously pigmented but not with a dark median
stripe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
54 Costal margins of both fore- and hindwings with a usually continuous
thick band of pigment extending to their tips (Fig. 96D). Thorax rather
pale except for conspicuous dark longitudinal lateral stripes (Myzocallis
subgenus Lineomyzocallis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
- Costal margins of wings without a continuous band of pigment. Thorax
variously pigmented but without contrasting dark lateral stripes . . . 68
55 R IV+V more than 0.10 mm long, longer than HT II . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis longirostris
- R IV+V usually less than 0.10 mm long and always shorter than HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
56 Hairs on ANT III finely pointed, 1-2 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis spinosa
- Hairs on ANT III with acute, blunt or slightly capitate apices, always
shorter than basal diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . 57
428 Quercus

57 ANT PT/BASE 3.5-4.5, BASE VI measuring less than 0.15 mm.

Immatures with long, pointed dorsal hairs . . . . . . . . Myzocallis elliotti
- ANT PT/BASE 1.0-3.5, with BASE VI more than 0.16 mm. Immatures
with at least some of dorsal hairs capitate . . . . . . . . . . . . . . . . . . . . . . 58
58 Fore tibiae distinctly darker than mid- and hind tibiae . . . . . . . . . . . 59
- All tibiae similarly pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
59 ANT PT/BASE 2.5-3.5. ANT III dusky on sensoriated section (use hand
lens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis longiunguis
- ANT PT/BASE 1.0-2.5. ANT III pale except on distal part . . . . . . 60
60 Fore femur with pigment dorsally and ventrally, at least on distal half
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis exultans
Fore femur unpigmented except sometimes at extreme apex . . . . . . 61
61 ANT PT/BASE 1.1-1.6. Pronotum with 3-5 posteriolateral hairs on each
side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis multisetis
- ANT PT/BASE 1.7-2.5. Pronotum usually with 2 posteriolateral hairs on
each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
62 Embryos with all spinal hairs short and not overlapping . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis occulta*
- Embryos with spinal hairs long and touching or overlapping . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis walshii
63 Pronotum with 1 posteriolateral hair on each side . . . . . . . . . . . . . . . 64
- Pronotum with 2 (or 3) posteriolateral hairs on each side . . . . . . . . 65
64 ANT IV and V uniformly dark. Immatures with 2 dorsal longitudinal rows
of dark spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis frisoni
- ANT IV and V pale basally shading to dark distally. Immatures without
dorsal pigmentation (except in ssp. iturbide) . . . . . . Myzocallis pepperi
65 Hind femur black on about distal third only . . . . . . . . . . . . . . . . . . . . 66
- Hind femur black except at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
66 Lobes of anal plate longer than their basal width, and CAUDA with an
elongate 'neck' (Fig. 96C) . . . . . . . . . . . . . . . . . . . . Myzocallis granovskyi
- CAUDA and anal plate normal for genus, i.e. lobes of anal plate shorter
than their basal width, and constricted part of CAUDA not elongated .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis durangoensis
67 ANT III-V usually pale except apically. ANT I and II often dark. Pro-
thoracic stripes 0.16-0.18 mm wide, with pale area between them longer
than wide (Fig. 96D). Dorsal hairs of immatures blunt or weakly capitate
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis bella
 Quercus 429

- ANT III-V usually wholly dusky, whereas ANT I and II are usually pale.
Prothoracic stripes 0.09-0.14 mm wide, with pale area between them at
least as wide as long (Fig. 96E) . . . . . . . . . . . . . . Myzocallis melanocera
68 PT 1.5 or more x ANT III; ANT PT/BASE 2.5-3.0. ANT III with 1-3
(usually 2) secondary rhinaria near base . . . . . . . . Myzocallis tenochca
- PT shorter than ANT III; ANT PT/BASE 1.2-3.2. ANT III with 2-12
(rarely 2) secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
69 Rs dark, with a black spot at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
- R s very pale, especially at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
70 R IV+V less than 0.11 mm, 0.80-0.95 x HT II . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis punctata (pale form)
- R IV+V 0.11 or more, 1.0-1.2 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis rostropunctata
71 Pterostigma of normal length, i.e. length measured from proximal end of
black patch 3.8-4.7 x maximum width (Fig. 96H) . . . . . . . . . . . . . . 72
- Pterostigma short, trapezoid, the length 2.5-3.8 x maximum width
(Fig. 96I) (subgenus Pasekia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
72 R IV+V stiletto-shaped, 0.15-0.22 mm long, 1.6-2.0 x HT II . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis occidentalis
- R IV+V 0.10-0.16 mm long, 0.8-1.5 x HT II . . . . . . . . . . . . . . . . . . 73
73 R IV+V 1.2-1.5 X HT II, with 4-7 accessory hairs . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis schreiberi
- R IV+V 0.84-1.19 x HT I I , with 4-12 accessory hairs . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis boerneri
74 ANT PT/BASE 2.4-3.2. (Spinal sclerites on ABD TERG 1.5 oval, with
longest hairs 28-50 µm long, having blunt or slightly capitate apices;
Fig. 96G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis cocciferina
- ANT PT/BASE 1.2-2.1. (Spinal sclerites on ABD TERG 1-5 and the
hairs upon them of various shapes and forms, but if longest hairs are more
than 40 µm long then they usually have strongly expanded or capitate
apices) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
75 Longest hairs on vertex very long, about 2.5 x basal diameter of ANT
III, and those on ANT III about equal to basal diameter of ANT III or a
little longer, and distinctly capitate. Primary rhinarium on ANT VI not
ciliated, elongate, 41-50µm long. ANT III with 8-12 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis persica
- Longest hairs on vertex 0.5-1.2 x basal diameter of ANT III, and those
on ANT III about 0.5 x basal diameter. Primary rhinarium on ANT VI
ciliated, less than 40 µm long. ANT III with 2-8 secondary rhinaria . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
430 Quercus

76 Empodial hairs setiform. Spinal hairs on ABD TERG 1-5 mostly pointed,
and all less than 40 µm long. Sclerites on ABD TERG 4 and 5 often fused
intersegmentally to form a pair of large dark patches . . . . . . . . . . . . . . .
.............................................. Myzocallis taurica
- Empodial hairs spatulate. Longest spinal hairs on ABD TERG 1-5 often
more than 40 µm long, with expanded or capitate apices. Sclerites on ABD
TERG 4 and 5 never fused intersegmentally . . . . . . . . . . . . . . . . . . . . 77
77 Hairs on vertex arising from pigmented sclerites (pale in spring forms).
ABD TERG 1-5 each with a pair of oval to quadrate sclerites that have
a dark margin around a large pale central area (Fig. 96F). Primary
rhinarium on ANT VI 26-33 µm long . . . . . . . . . . Myzocallis komareki
- Vertex without pigmented sclerites at hair-bases, but with paired longi-
tudinal bands of brown pigment running through lateral ocelli (indistinct
in spring forms). Paired spinal sclerites on ABD TERG 1-5 oval or
irregular in shape, often paler in centre but without a clearly defined dark
margin. Primary rhinarium on ANT VI 19-25 µm long . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis mediterraneus

KEY F - Drepanosiphinae and Thelaxinae with apterous viviparae on oaks

Refers to apterous viviparae only except where otherwise stated.
1 Eyes of 3 facets. Antennae 5-segmented. Dorsal hairs fine if long, or short
and thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Eyes multifaceted. Antennae 4- to 6-segmented; if 4- or 5-segmented then
dorsal hairs very long and thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2 CAUDA rounded. Dorsal hairs all long and fine . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kurisakia querciphila
- CAUDA knobbed. Many dorsal hairs short, thick and spine-like, dagger-
like or club-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 ABD TERG 5 with 12-25 short, blunt, club-shaped hairs (Fig. 97A). Head
and pronotum with 50-67 and mesonotum with 30-53 hairs . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelaxes valtadorosi
- ABD TERG 5 with 8-14 pointed, spine-like (Fig. 97B) or dagger-like
(Fig. 97C) hairs. Head and pronotum with 20-26 hairs, mesonotum with
13-28 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 R V less than 0.25 x R IV and less than 2 x ANT PT. Spinal hairs on
ABD TERG 5 usually less than 24 µm long, spine-like (Fig. 97B). (Alata
with 4-7 secondary rhinaria on ANT III) . . . . . . . . Thelaxes dryophila
- R V more than 0.25 x R IV and 2 or more x ANT PT. Spinal hairs on
ABD TERG 5 mostly more than 25 µm long, very thick and dagger-like
(Fig. 97C). (Alata with 0-5 secondary rhinaria on ANT III) . . . . . . . 5
5 ANT PT/BASE 0.17-0.24. Knob of CAUDA slightly broader than long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelaxes suberi
 Quercus 431

Fig. 97. Spinal hairs on ABD TERG 5 of apterae of A, Thelaxes valtadorosi, B, Th. dryophila,
C, Th. suberi, D, Siculaphis vittoriensis, E, Hoplochaitophorus heterotrichus, f, H. quercicola; G, CAUDA
of Globulicaudaphis pakistanica (ventral view).

- ANT PT/BASE 0.26-0.40. Knob of CAUDA at least as long as broad

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelaxes californica
6 Dorsal hairs mostly conspicuously bifurcate, resembling fishes' tails
(Fig. 97D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siculaphis vittoriensis
- Dorsal hairs not bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 At least some of dorsal hairs thick and spine-like or capitate, arising from
tuberculate bases or from elongate processes . . . . . . . . . . . . . . . . . . . . . 8
- Dorsal hairs all thin and setaceous, sometimes with expanded apices but
not arising from tuberculate bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
432 Quercus

8 Antennae 4- or 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Antennae 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
9 Dorsal cuticle strongly and densely nodulose. Frons with 2 long forwardly-
directed hair-bearing processes, and thorax and abdomen with large
lateral processes bearing backwardly-directed hairs . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis areolatus
- Dorsal cuticle spiculose, wrinkled or very weakly nodulose. Frons without
processes, and thorax and abdomen with marginal hairs on tuberculate
bases not much longer than their basal diameters . . . . . . . . . . . . . . . 10
10 ANT PT/BASE 0.6-0.7. Dorsal hairs on head and pronotum less than
20 µm long. Dorsal cuticle with faint nodulose sculpturing . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis calvus
- ANT PT/BASE 0.7-1.4. Dorsal hairs on head and pronotum longer than
25 µm. Dorsal cuticle spiculose or wrinkled . . . . . . . . . . . . . . . . . . . . . 11
11 Dorsal cuticle spiculose. Antennae of 4 or 5 segments, with ANT III only
1-2 x SIPH. SIPH 0.046-0.166 mm long, 2-6 x their minimum diameter
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis analiliae
Dorsal cuticle not spiculose. Antennae always 4-segmented, with ANT III
3.8-6.8 x SIPH. S1PH 0.027-0.044 mm long, less than twice their mini-
mum diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mexicallis spinifer
12 CAUDA bluntly conical, without a constriction, anal plate only weakly
indented. Dorsal spine-like hairs bearing many minute spinules . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Serralocallis takahashii
- CAUDA knobbed, anal plate bilobed. Dorsal spine-like hairs not bearing
minute spicules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Spine-like dorsal hairs outnumbered by fine hairs with finely-pointed
apices (Fig. 97E; sometimes spine-like hairs are few in number, or even
absent in weakly-pigmented midsummer individuals). Longest hairs on
ANT III long and fine, at least twice basal diameter of segment . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplochaitophorus heterotrichus
- Most dorsal body hairs spine-like and on tuberculate bases (e.g. Fig. 97F)
Longest hairs on ANT III 1.0-1.25 x basal diameter of segment . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 SIPH flared apically and almost as long as R IV+V. ANT PT/BASE more
than 0.65 . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplochaitophorus quercicola
- SIPH small, not much widened apically, about 0.5 x R IV+V. ANT
PT/BASE less than 0.60 . . . . . . . . . . . . . Hoplochaitophorus spiniferus
15 Anal plate bilobed, CAUDA with or without a constriction. Wax pore
plates not evident. ANT II shorter than ANT I . . . . . . . . . . . . . . . . . 16
 Quercus 433

- Anal plate entire, CAUDA broadly rounded. Wax pore plates evident.
ANT II much longer than ANT I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
16 ANT PT/BASE 1.0 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- ANT PT/BASE more than 1.2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
17 Many of dorsal body hairs very long, up to or exceeding length of ANT
III. CAUDA knobbed . . . . . . . . . . . . . . . . Hoplochaetaphis zachvatkini
- Dorsal body hairs all very much shorter than ANT III. CAUDA with or
without a constriction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 Antennae densely hairy; ANT VI BASE with 7-15 hairs. (Alata with
forewing veins thickly bordered with fuscous) . . . . . . . . . . . . . . . . . . . 19
- Antennae sparsely hairy; ANT VI BASE with 1-3 hairs. (Alata with
forewing veins not or only weakly bordered) . . . . . . . . . . . . . . . . . . . . 21
19 ANT PT/BASE about 0.6 . . . . . . . . . . . . . . . . . . . . . . . Patchia winforii*
- ANT PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Alata with 4-8 (most commonly 5 or 6) secondary rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnochaitophorus querceus
- Alata with 6-11 (most commonly 8 or 9) secondary rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnochaitophorus obscurus
21 Mid-femora, hind femora and hind tibiae mainly dark . . . . . . . . . . . 22
- Mid-femora, hind femora and tibiae mainly pale . . . . . . . . . . . . . . . . 25
22 ABD TERG 8 with more than 25 hairs . . . . . . . . . . . . . . . . . . . . . . . . 23
- ABD TERG 8 with less than 16 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 24
23 ANT PT/BASE less than 0.4. ABD TERG 1-5 without transverse bars,
but with small sclerites at bases of the very short dorsal hairs. R IV+V
with 8-11 accessory hairs . . . . . . . . . . . . . . . . Neosymydobius agrifoliae
- ANT PT/BASE more than 0.5. ABD TERG 1-5 with transverse bars and
long fine hairs (more than 50 µm). R IV+V with 4-6 accessory hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius albasiphus/canadensis
24 Hairs on ABD TERG 1-6 pointed, blunt or very slightly capitate. Hairs
on ANT III pointed, more than 0.5 x basal diameter of segment . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius chrysolepis
- Hairs on ABD TERG 1-6 flared apically. Hairs on ANT III blunt, less
than 0.5 x basal diameter of segment . . . . . . . Neosymydobius butzei
25 ABD TERG 1-7 each with 1-3 (usually 2) marginal hairs on each side
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
- ABD TERG 1-7 each with 3-7 marginal hairs on each side . . . . . . 27
434 Quercus

26 Marginal tubercles present on all tergites (10-12 on pronotum, 1-4 on

other segments). ANT PT/BASE less than 0.6. ABD TERG 1-6 with
broad but only lightly pigmented transverse bands each bearing 12-16
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius ajuscanus
- Marginal tubercles absent or rarely present. ANT PT/BASE more than
0.6. ABD TERG 1-6 with well-pigmented transverse bands bearing only
6 hairs (2 spinal, 4 marginal) . . . . . . . . . . . . . . . Neosymydobius mimicus
27 Hairs on ANT III long and fine, 2-3 x basal diameter of segment. ANT
PT/BASE less than 0.8 . . . . . . . . . . . . . . Neosymydobius quercihabitus
- Hairs on ANT III shorter than basal diameter of segment. ANT PT/
BASE more than 0.8 . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius luteus
28 Dorsal abdominal hairs very short, pointed. Marginal tubercles present on
all tergites (5-10 on pronotum, 0-5 on other segments) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosymydobius memorialis
- Dorsal abdominal hairs long with blunt, swollen, capitate or flared apices.
Marginal tubercles absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29 CAUDA not knobbed, but swollen dorsally into a membranous, hemi-
spherical bladder with a sclerotic, hair-bearing plate on the underside
(Fig. 97G) . . . . . . . . . . . . . . . . . . . . . . . . . . . Globulicaudaphis pakistanica
- CAUDA knobbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
30 Spinopleural hairs on ABD TERG 1-6 very numerous (more than 30 per
segment), 15-55µm long. Longest hairs on ANT III strongly capitate,
longer than basal diameter of segment . . . . . . . . Myzocallis polychaeta
- ABD TERG 1-6 each with 8-14 spinopleural hairs 15-140 µm long.
Longest hairs on ANT III blunt or weakly capitate, shorter than basal
diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31 Antennae banded, tibiae mainly pale. ABD TERG 1-7 usually with dark
spinal sclerites, not fused between segments. ANT PT/BASE 1.4-1.7.
(Forewing of alata unpigmented except for small dark spots on the
pterostigma and at base of Cu lb ) . . . . . . . . . . . . . Myzocallis glandulosa
- Antennae and tibiae mainly dark. Dorsal abdomen with a black spinal
patch restricted to ABD TERG 3-5, fused between segments. ANT PT/
BASE 2.3-3.0. (Alata with entire costal margin of forewing pigmented)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzocallis meridionalis
32 SIPH pores small to minute (less than 25 µm in diameter), not surrounded
by a ring of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
- SIPH pores larger (more than 25 µm in diameter), with an encircling ring
of hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
33 SIPH pore much larger than (4-6 x diameter of) nearest wax pores. Fore
femur 4-5 x its maximum width. R IV+V with convex sides, somewhat
pointed apically but not stiletto-shaped (e.g. Fig. 98A) . . . . . . . . . . . 34
 Quercus 435

Fig. 98. R IV+V of A, Diphyllaphis quercus, B, D. konarae, C, D. microtrema, D, Stegopylla quercicola.

- SIPH pore only 1-2 x diameter of nearest wax pores. Fore femur 2.5-3.5
x its maximum width. R IV+V with slightly concave sides, either blunt
apically or stiletto-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
34 ABD TERG 8 with 2 hairs. ANT PT/BASE 0.2 or less . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis mordvilkoi
- ABD TERG 8 with 4 hairs. ANT PT/BASE more than 0.3 . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis quercus
35 ABD TERG 8 with 8-10 hairs, CAUDA with 2 hairs. R IV+V blunt
apically (Fig. 98C), with primary (subapical) hairs not displaced proxi-
mally (all on distal third) . . . . . . . . . . . . . . . . . Diphyllaphis microtrema
- ABD TERG 8 with 4 hairs, CAUDA with 3-4 hairs. R IV+V pointed or
stiletto-shaped, with primary hairs displaced proximally (Fig. 98B) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
36 ANT II shorter than ANT III. R IV+V shorter than HT II. ABD TERG
6-8 fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis konarae
- ANT II longer than ANT III. R IV+V longer than HT II. ABD TERG
6-8 with separate sclerites . . . . . . . . . . . . . . . . . . . . . . . Diphyllaphis alba
37 Second tarsal segments with lateral and often also ventral hairs at about
midlength, in addition to the subapical hairs . . . . . . . . . . . . . . . . . . . . 38
- Second tarsal segments with only subapical hairs . . . . . . . . . . . . . . . . 39
38 R IV+V 0.7-0.85 x HT II, with one pair of primary hairs displaced
proximally (in addition to a pair of accessory hairs; Fig. 98D) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla quercicola
- R IV+V 0,95-1.05 x HT II, with subapical hairs not displaced . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla mugnozae
39 R IV+V 0.13-0.15 mm long, 1.2 or more x HT II . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla quercifoliae
- R IV+V 0.07-0.10 mm long, 0.7-1.1 x HT II . . . . . . . . . . . . . . . . . . 40
436 Radermachera

40 R IV+V 0.07-0.08 mm long, 1.0-1.1 x ANT II and 0.7-0.9 x HT II

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla querci
- R IV+V 0.085-0.10 mm long, 1.2-1.6 x ANT II and 0.9-1.1 X HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegophylla essigi

RADERMACHERA Bignoniaceae

Myzus persicae

RANDIA Rubiaceae

Randia sinensis Toxoptera aurantii

R. spinosa Myzus persicae
Toxoptera aurantii
Randia sp. Taiwanaphis randiae

Key to aphids on Randia

- ANT PT/BASE about 1. SIPH short, dark, conical, much shorter than
their basal width. CAUDA knobbed, anal plate bilobed. ANT III of alata
with 32-46 transversely oval secondary rhinaria, not in a row . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis randiae
- ANT PT/BASE more than 2. SIPH tubular. CAUDA tongue-shaped,
anal plate entire. ANT III of alata with 2-17 roundish secondary rhinaria
in a single row . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

RAPANEA Myrsinaceae

Rapanea spp. Toxoptera aurantii

 Rhamnus 437

RAPHIA Palmae

Cerataphis variabilis

RAUWOLFIA Apocynaceae

Host Plant List

Rauwolfia caffra Sitobion halli
R. inebrians Sitobion halli
R. lamarkii Aphis spiraecola
R. tetraphylla Aphis spiraecola
R, vomitoria Sitobion halli, krahi

Key to aphids on Rauwolfia

1 Antennal tubercles not developed. ANT PT/BASE less than 3. SIPH
without any subapical reticulation. CAUDA black, with a slight mid-way
constriction (Fig. 121T) . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
Antennal tubercles well developed, divergent. ANT PT/BASE more than
6. SIPH with a subapical zone of polygonal reticulation. CAUDA pale,
without a constriction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Longest posterior dorsal cephalic hairs (between eyes) less than 18µm
long. R IV+V 0.80-1.05 x HT II. SIPH narrow (about 0.5 x width of
CAUDA at their respective midpoints), with zone of polygonal reticula-
tion extending less than 0.2 of its total length . . . . . . . . . Sitobion halli
- Longest posterior cephalic hairs more than 18µm long. R IV+V
1.05-1.20 x HT II. SIPH broad (about as wide as CAUDA at their
respective midpoints), with zone of polygonal reticulation extending more
than 0.3 of total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion krahi

RHAMNUS Rhamnaceae

Host Plant List

Rhamnus alaternus Aphis craccivora, frangulae group,
nasturtii
438 Rhamnus

Macchiatiella rhamni
Toxoptera aurantii
Rh. alnifoliae Aphis nasturtii
Rh. alpina Macchiaüella rhamni
Rh. californica Sitobion rhamni
Rh. cathartica Aphis commensalis, frangulae
group, gossypii, mammulata,
nasturtii, spiraecola
Macchiatiella rhamni, rhamni ssp.
tarani
Rh. dahurica Aphis glycines, gossypii
Rh. frangula Aphis frangulae group, gossypii,
nasturtii
Macchiatiella rhamni
Rh. japonica Aphis gossypii
Macchiatiella itadori
Rh. koraiensis Aphis gossypii
Rh. lanceolata Macchiatiella rhamni
Rh. libanoticus Aphis craccivora, frangulae group,
spiraecola
Rh. nepalensis Myzus ornatus
Rh. palaestinus Aphis craccivora, frangulae group,
spiraecola
Rh. purshiana Aphis gossypii
Macchiatiella itadori, rhamni
[Myzus lythri]
Sitobion rhamni
Rh. utilis Aphis utilis
Rh. virgatus Aphis rhamnifila
Rhamnus sp. Aphis fabae group
Myzus persicae

Key to Rhamnus-feeding species (apterae viviparae except fundatrices)

1 CAUDA very broadly rounded, less than 0.5 x basal width. Dorsal
abdomen usually with distinctive black markings . . . . . . . . . . . . . . . . . 2
- CAUDA helmet-, tongue- or finger-shaped, 0.6 or more x its basal width.
Dorsal abdomen with or without dark markings . . . . . . . . . . . . . . . . . . 3
2 SIPH black. Dorsal abdomen with an extensive solid black patch.
R IV+V 1.15-1.28 x HT II . . . . . . . . . . . . . . . . . . Macchiatiella itadori
 Rhamnus 439

- SIPH pale. Dorsal abdomen with dark patch broken centrally or in

midline, often on ABD TERG 3-4 (-5) only and sometimes absent.
R IV+V 1.0-1.15 x HT II . . . . . . . . . . . . . . . . . . . Macchiatiella rhamni
3 Head spiculose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- Head smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4 ANT III with a small secondary rhinarium near base. Antennal tubercles
with inner faces almost parallel. R IV+V 1.25-1.40 x HT II. SIPH
straight, tapering, dark at least on distal half. CAUDA also dark. (Alata
with dark transverse bars on dorsal abdomen) . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum rhamni
- ANT I I I without any rhinaria. Antennal tubercles strongly convergent.
R IV+V 0.09-1.18 x HT I I . SIPH slightly swollen or curved outwards
distally. SIPH and CAUDA mainly pale. (Alata with a black dorsal
abdominal patch) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Dorsal abdomen with an intersegmental pattern of dark spots and flecks.
ANT PT/BASE 2.5 or less. SIPH tapering with shallow S-curve, not
swollen distally (Fig. 121J) . . . . . . . . . . . . . . . . . . . . . . . . . Myzus ornatus
- Dorsal abdomen without dark intersegmental markings. ANT PT/BASE
more than 3. SIPH slightly swollen on distal half (Fig. 121H) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
6 Antennal tubercles well developed, divergent. ANT PT/BASE more than
6. ANT I I I with 1-2 secondary rhinaria near base. SIPH pale, with
subapical polygonal reticulation, 0.3 or more x BL . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sitobion rhamni
- Antennal tubercles weakly developed. ANT III without rhinaria. ANT
PT/BASE less than 5. SIPH pale or dark, without subapical polygonal
reticulation, less than 0.2 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ANT PT/BASE more than 3.5. Stridulatory ridges present on abdominal
sternites 5 and 6, and a row of peg-like hairs on hind tibia (Fig. 122)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- ANT PT/BASE less than 3.5. No Stridulatory apparatus . . . . . . . . . . 8
8 Eongest hairs on ANT III 1.1-2.4 x basal diameter of segment . . . 9
- Longest hairs on ANT III 0.4-0.8 x basal diameter of segment . . . . 10
9 Pronotum and ABD TERG 1-4 and 7 with extremely large, hemispherical
marginal tubercles. SIPH 1.5-2.0 x CAUDA and 4-7 x their width at
midpoint. Antennae 6-segmented . . . . . . . . . . . . . . . . Aphis mammulata
- Marginal tubercles of normal size on ABD TERG 1 and 7, and absent
from ABD TERG 2-4. SIPH 0.9-1.2 x CAUDA, and 1.5-3 x their width
at midpoint. Antennae 5- or 6-segmented . . . . . . . . . Aphis commensalis
440 Rhamnus

10 ABD TERG 1-4 and 7 with large, hemispherical marginal tubercles

diameter of those on ABD TERG 2-4 equal to or greater than midlength
diameter of SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis versicolor
- Marginal tubercles of normal size on ABD TERG 1 and 7, and small or
absent on ABD TERG 2-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 SIPH and CAUDA both pale, or SIPH dark only towards apices. (Alata
with 8-18 secondary rhinaria on ANT III, 1-6 on IV and 0-2 on V
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis nasturtii
- SIPH dark, CAUDA pale or dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 Dorsal abdomen almost wholly covered by an extensive dark sclerite
CAUDA black and usually with 4-7 hairs (Fig. 121N) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
- Dorsal abdomen without an extensive solid dark sclerite. CAUDA if black
then usually with more hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.
13 CAUDA dark with 11-24 hairs (rarely less than 12). Femora dark at least
distally. ABD TERG 7 and 8 with dark cross-bands, and more anterior
tergites often with some dark markings (Fig. 121R) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis fabae group
- CAUDA pale or dark, with 4-15 hairs (rarely more than 12). Femora pale
or dusky. Dorsal abdomen without dark markings . . . . . . . . . . . . . . . 14
14 CAUDA as dark as SIPH with 7-15 (usually 8-12) hairs and a slight con-
striction at or just proximal to midpoint (Fig. 121T) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
- CAUDA paler than SIPH, with or without a constriction, or if as dark
as SIPH then with only 4-7 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Dorsal cephalic hairs finely pointed, 35-45 µm long. Hind femur with
some ventral hairs over 45 µm long, exceeding diameter of trochantro-
femoral suture. CAUDA almost as dark as SIPH, with about 6 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis rhamnifila
- Dorsal cephalic hairs blunt, less than 30 µm long. No hairs on hind femur
exceeding diameter of trochantro-femoral suture. CAUDA usually paler
than SIPH, with 4-10 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 Head and ANT I-V pale, only ANT VI dusky. SIPH usually more than
0.2 X BL. CAUDA much paler than SIPH, often with a midpoint con-
striction, and bearing 8-10 hairs . . . . . . . . . . . . . . . . . . . . Aphis glycines
- Head and ANT I-VI dusky. SIPH less than 0.2 x BL. CAUDA often
dusky, not contrasting with SIPH, unconstricted, bearing 4-9 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis frangulae group
 Rhus 441

RHAPIS Palmaceae

Rhapis javanicus Astegopteryx rhapidis

(For palm aphids see B & E, 1984)

RHUS Sumacs Anacardiaceae

A revised and extended version of the key given in Blackman and Eastop
(1984) follows the host plant list. Subgenus Toxicodendron, which does not
contain any trees, is not included.

Host Plant List

Rhus abyssinica Aphis rhoicola
Rh. ambigua Carolinaia floris
Rh. aromatica Carolinaia rhois
Rh. crenata Aphis gossypii
Rh. dentata Aphis gossypii
Rh. glabra Carolinaia rhois
(Smooth Sumac) Melaphis rhois
Rh. javanica = Rh. semialata
Rh. laevigata Aphis gossypii
Rh. lancea Aphis gossypii
Myzus persicae
Rh. lucida Aphis gossypii
Rh. potanini Kaburagia ovatirhusicola,
rhusicola
[Nurudea choui]
Schlechtendalia microgallis
Rh. punjabensis var. sinica Kaburagia rhusicola
Nurudea meitanensis
Schlechtendalia elongallis
Rh. pyroides Aphis gossypii
Rh. semialata [Aphis craccivora]
(Chinese Sumac) Chuansicallis chengtuensis
442 Rhus

[Macrosiphum euphorbiae]
Nurudea ibofushi, shiraii,
yanoniella
[Prociphilus oriens]
Schlechtendalia chinensis
Toxoptera odinae
Rh. succedanea Carolinaia japonica
(Japanese Sumac, Wax-tree) Toxoptera odinae
Rh. trichocarpa Carolinaia japonica, nigra
Rh. typhina Carolinaia rhois
(Staghorn Sumac) Melaphis rhois
Rh. vernicifera Carolinaia rhois
(Varnish Tree) Stomaphis rhusivermiciflua
Rh. viminalis Aphis gossypii
Myzus persicae
Rhus spp. indet. Aphis gossypii
Aulacorthum magnoliae

Key to aphids on Rhus

1 Very large aphid, BL about 5 rnm or more, with very long rostrum about
1.5 x BL . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis rhusivermiciflua
- BL less than 3 mm, rostrum much shorter than BL . . . . . . . . . . . . . . 2
2 ANT PT/BASE more than 0.5, with PT finger-like. SIPH present . ..
............................................................. 3
- ANT PT/BASE much less than 0.5, with PT short and thick. SIPH absent
(gall formers) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3 ANT PT/BASE less than 1. SIPH as small, truncate cones, shorter than
basal width. CAUDA knobbed, anal plate bilobed . . . . . . . . . . . . . . . . .
....................................... Chuansicallis chengtuensis
- ANT PT/BASE more than 1. SIPH tubular, longer than basal width.
CAUDA tongue- or finger-like, anal plate entire . . . . . . . . . . . . . . . . . 4
4 SIPH shorter than CAUDA. Stridulatory ridges present ventrolaterally on
abdominal sternites 5 and 6 . . . . . . . . . . . . . . . . . . . . . Toxoptera odinae
- SIPH longer than CAUDA. No Stridulatory ridges . . . . . . . . . . . . . . . 5
5 SIPH tapering continuously from base to flange . . . . . . . . . . . . . . . . . 6
- SIPH swollen at least slightly in middle or on distal half . . . . . . . . . 7
6 ANT PT/BASE less than 2 . . . . . . . . . . . . . . . . . . . . . . . . Aphis rhoicola
- ANT PT/BASE more than 2 . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
 Rhus 443

7 Head capsule smooth or wrinkled. ANT I I I (distally) to VI, tibiae and

SIPH dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glabromyzus rhois
- Head capsule scabrous or spiculose, at least ventrally. Antennae, tibiae
and SIPH pale or dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8 Antennal tubercles low and divergent. SIPH dark . . . . . . . . . . . . . . . . 9
- Antennal tubercles well developed, with inner faces parallel or convergent.
SIPH pale, except at apex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9 Femora strongly scabrous and spiculose distally for more than half of
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carolinaia niger
- Femora smooth or imbricated distally, but without spicules on the
imbrications . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 ANT PT/BASE 1.5-2.0. R 1V+V 1.0-1.1 x HT I I . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carolinaia floris
- ANT PT/BASE 2.1-3.0. R IV+V 1.2-1.4 x HT II . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carolinaia rhois
11 Antennae and legs mainly dark. ANT III with 1-2 secondary rhinaria near
base. Antennal tubercles with inner faces parallel . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum magnoliae
- Antennae and legs mainly pale. ANT III without any rhinaria. Antennal
tubercles with inner faces convergent . . . . . . . . . . . . . . . Myzus persicae
12 (Remaining couplets refer to alate migrants from galls) Pterostigma of
forewing extending in a curve around tip of wing (Fig. 99A) . . . . . . 13
- Pterostigma of normal length, not extending around wing-tip (Fig. 99B)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13 Antenna 6-segmented, II-VI each with a single very large rhinarium
occupying most of its surface area, in which are scattered many small
islands of sclerotized cuticle (Fig. 99C) . . . . . Schlechtendalia elongallis
- Antenna 5-segmented; II-V each with at least partial separation of the sen-
soriated surface into smaller units (Fig. 99D) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schlechtendalia chinensis
14 Antenna 6-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- Antenna 5-segmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
15 ANT III-VI each with a single large oblong rhinarium occupying most of
its surface area (Fig. 99E) . . . . . . . . . . . . . . . . . . . . . Kaburagia rhusicola
- ANT III-VI with separate, transversely elongate rhinaria . . . . . . . . . 16
16 Antenna with secondary rhinaria distributed ANT III 6-13, IV 3-9,
V 3-11, VI 5-11 (Fig. 99F) . . . . . . . . . . . . . . . . . . . Melaphis rhois (part)
444 Rhus
 Robinia 445

- Antenna with secondary rhinaria distributed ANT I I I 2-5, IV 1-4, V 2-4,

VI 3-4 (Fig. 99G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nurudea meitanensis
17 Antenna with secondary rhinaria distributed ANT III 9-24, IV 3-9,
V 5-11 (Fig. 99H) . . . . . . . . . . . . . . . . . . . . . . . . . . . Melaphis rhois (part)
- Antenna with secondary rhinaria distributed ANT III 3-11, IV 1-7, V 1-9
(Figs 99I-K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18 ANT IV and V each with a single large rhinarium occupying most of the
surface area, enclosing small scattered islands of cuticle (Fig. 99I) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nurudea ibofushi
- ANT IV and V with separated, annular rhinaria . . . . . . . . . . . . . . . . 19
19 Rhinaria on ANT IV and V mostly broad and strap-like, and sometimes
joined to form rhinaria longer than width of antenna (Fig. 99J) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nurudea shiraii
- Rhinaria on ANT IV and V all narrow and separate (Fig. 99K) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nurudea yanoniella

RICINUS Euphorbiaceae

Ricinus communis Aphis gossypii

(Castor Oil Plant) Myzus persicae
(Use key to polyphagous aphids, p. 532)

ROBINIA Leguminosae

Robinia neomexicana Appendiseta robiniae

R. pseudoacacia Acyrthosiphon gossypii, pisum
(Black Locust) Aphis craccivora, craccivora ssp.
pseudoacaciae, fabae, gossypii,
spiraecola

Fig. 99. Forewing of A, Schlechtendalia chinensis and B, Kaburagia rhusicola; antenna of alata from gall
of C, Schlechtendalia elongallis (from Tsai and Tang, 1946), D, S. chinensis, E, Kaburagia rhusicola
(from Tsai and Tang, 1946, as Macrorhinarium ovogallis), F, Melaphis rhois (6-segmented), G, Nurudea
meitanensis (from Tsai and Tang, 1946, as Floraphis), H, Melaphis rhois (5-segmented), I, Nurudea
ibofushi, J, N. shiraii, K, N. yanoniella.
446 Rothmannia

Appendiseta robiniae
Myzus persicae
[Protrama orientalis Narzikulov,
1962]
[ Tinocallis zelkowae]

Key to aphids on Robinia

1 All viviparae alate, with CAUDA knobbed, anal plate bilobed. Secondary
rhinaria on ANT III transversely elongate. ANT PT/BASE about 0.5.
SIPH short, truncated cones with a single short hair attached at base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendiseta robiniae
Apterous viviparae normally present. CAUDA tongue- or finger-shaped,
anal plate entire. Secondary rhinaria round or oval. ANT PT/BASE more
than 2. SIPH tubular, without attached hair . . . . . . . . . . . . . . . . . . . . . 2
2 Antennal tubercles well developed, broadly divergent, smooth. SIPH
long, pale, tapering from a broad base, without subapical reticulation
............................................................. 3
Antennal tubercles weakly developed or, if well developed, convergent
and scabrous. SIPH dark or pale, if pale then slightly swollen on distal
half or with subapical reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
3 SIPH 0.33-0.50 x BL and 2.5-3.5 x CAUDA . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon gossypii
- SIPH 0.20-0.36 x BL and 1.2-1.8 x CAUDA . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon pisum

ROTHMANNIA Rubiaceae

Rothmannia capensis Toxoptera citricidus

SALACCA Palmae

Salacca edulis Astegopteryx nipae

Cerataphis variabilis
(For palm aphids see B & E, 1984)
 Salix 447

SALIX Willows Salicaceae

Over 120 aphid species feed on Salix, the main groups belonging to the genus
Chaitophorus (48 species), the Pterocommatini (29 species in 4 genera) and
the genus Cavariella (20 species), the latter having host alternation between
Salix and Umbellifereae or Araliaceae. Although almost all willow-feeding
aphids restrict themselves exclusively to the genus Salix there are no clear
patterns of host specificity within the genus. Therefore, following the host
plant list, the aphids are keyed for the genus as a whole; the three main groups
noted above are taken out at couplets 16-17 and keyed separately in sup-
plementary keys A, B and C respectively.

Host Plant List

Salix abscondita Chaitophorus horii

S. acmophylla Chaitophorus pakistanicus
(Bada, Bisu, Gadhbhains) Tuberolachnus salignus
S. acutifolia Aphis farinosa
(Caspic Willow) Cavariella konoi
Chaitophorus diversisetosus, niger,
quinquemaculatus, ramicola
Pterocomma konoi
S. aegyptiaca Tuberolachnus salignus
S. agrophylla Cavariella pustula
Chaitophorus macrostychae
S. alba (incl. var. coerulea, var. Aphis farinosa
sericea) Cavariella aegopodii,
(Pil, Vitpil, White Willow) archangelicae, nipponica,
paslinacae, salicicola, salicis,
theobaldi
Chaiiophorus israeliticus, niger,
nigritus, viminalis, vitellinae,
vitellinae ssp. danubicus
Neopterocomma asiphum
Phylloxerina salicis
Plocamaphis amerinae
Pterocomma bicolor,
chaetosiphon, konoi, pilosum,
pilosum ssp. sarmaticum,
salicis, smithiae, ringdahli,
rufipes
448 Salix

Stomaphis longirostris
Tuberolachnus salignus
S. amygdaloides Chaitophorus pusillus
(Peachleaf Willow) Macrosiphum californicum
S. apennina Chaitophorus salicti
S. arbuscula Cavariella intermedia
S. arbustifolia Cavariella salicicola
S. arctica Cavariella borealis
S. aurita Aphis farinosa
Cavariella intermedia
Chaitophorus capreae, ramicola,
salicti
Plocamaphis flocculosa
Pterocomma jacksoni, konoi,
pilosum
S. babylonica Aphis farinosa, gossypii
(Weeping Willow) Cavariella aegopodii, aquatica,
bonica, indica, konoi,
pastinacae, saliapterus,
salicicola, salicis
Chaitophorus beuthani, niger,
pakistanicus, salijaponicus,
saliniger, truncatus, viminalis,
vitellinae
Lachnus salicis
Phylloxerina salicis
Plocamaphis salijaponica
Pterocomrna bicolor, pilosum,
salicis, [salijaponica],
sanguiceps
Toxoptera aurantii
Tuberolachnus salignus
S. bakko Chaitophorus matsumurai
Prociphilus oriens
S. bebbiana Chaitophorus horii, saliapterus
Pterocomma konoi, rufipes
S. brachypoda Chaitophorus niger
S. caesia Chaitophorus truncatus
S. canariensis Tuberolachnus salignus
S. caprea Aphis farinosa
(Goat Willow, Sallow) Cavariella aegopodii,
 Salix 449

archangelicae, nipponica,
pastinacae, theobaldi
Chaitophorus capreae, flavissimus,
furcatus, horii, niger,
quinquemaculatus, ramicola,
salicti, salijaponicus, vitellinae
[Lachnus distinguendus Dahlbom,
1851]
Pachypappa warschavensis
Phylloxerina capreae
Plocamaphis flocculosa (sspp.
brachysiphon, goernitzi)
Pterocomma jacksoni, konoi,
pilosum, rufipes, salicis
S. cardiophylla Pterocomma rufipes
S. caroliniana Chaitophorus longipes, minutus
Phylloxerina sp.
Tuberolachnus salignus
S. chaenomeloides Phylloxerina capreae
Tuberolachnus salignus
S. chrysostella Cavariella aegopodii
S. cinerea Aphis farinosa
(Grey Willow) Cavariella aegopodii, japonica,
theobaldi
Chaitophorus capreae, niger,
ramicola, salicti
Plocamaphis flocculosa (sspp.
brachysiphon, goernitzi)
Pterocomma jacksoni, konoi,
pilosum, rufipes, salicis
Tuberolachnus salignus
S. cordata Chaitophorus nigrae, viminalis
S. crassijulis Plocamaphis coreana
S. daphnoides Aphis farinosa (incl. ssp.
(Violet Willow) yanagicola)
Cavariella aegopodii
Chaitophorus salicti
Phylloxerina daphnoides
Pterocomma pilosum
Tuberolachnus salignus
S. dasyclados = S. smithiana
S. decipiens Cavariella konoi
450 Salix

S. denticulata = S. elegans
S. dichroa Cavariella archangelicae
S. discolor Chaitophorus pallipes, pusillus
(American Pussy Willow) Phylloxerina salicicola
Plocamaphis flocculosa
Tuberolachnus salignus
S. dshugdshurica Chaitophorus saliapterus
Pterocomma rufipes
S. eleagnos = S. incana
S. elegans Cavariella aegopodii, biswasi,
nigrae, salicicola
Neoacyrthosiphon dubium
S. elegantissima Chaitophorus niger
Tuberolachnus salignus
S. eriocarpa Cavariella salicicola
Chaitophorus salijaponicus
S. exigua Chaitophorus macrostachyae,
(Sandbar Willow) viminalis
Pterocomma sanguiceps
S. fluviatilis Chaitophorus nigrae, populicola
(River Willow) Tuberolachnus salignus
S. fragilis Aphis farinosa
(Crack Willow) Cavariella aegopodii,
archangelicae, konoi,
pastinacae, quinquemaculalus,
theobaldi
Chaitophorus beulhani, niger,
truncatus, vitellinae
Lachnus longirostrum
Neopterocomma asiphum
Pemphigus saliciradicis
Phylloxerina salicis
Pterocomma chaetosiphon, konoi,
pilosum, salicis, smithiae
Tuberolachnus salignus
S. fulvopubescens Cavariella ?aegopodii, japonica
S. fuscescens Pterocomma rufipes
S. gilgiana Cavariella salicicola
S. glandulosa (var. glabra) Aphis farinosa
Cavariella salicicola
 Salix 451

Chaitophorus saliciniger
Plocamaphis coreana
Tuberolachnus salignus
S. glauca Cavariella konoi
Chaitophorus lapponum
Pterocomma smithiae
S. graciliglans Chaitophorus saliciniger
Plocamaphis coreana
S. gracilistyla Aphis farinosa
Cavariella archangelicae,
nipponica
Chaitophorus niger, saliciniger
Pterocomma rufipes, [salijaponica]
Tuberolachnus salignus
S. hastata Aphis farinosa
S. herbacea Cavariella aegopodii
Pemphigus saliciradicis
S. humilis Phylloxerina salicicola
S. incana Chaitophorus salicti, similis
Phylloxerina daphnoides
Tuberolachnus salignus
S. integra Cavariella aegopodii, nipponica
Chaitophorus matsumurai,
saliapterus, salijaponicus
S. irrorata Cavariella archangelicae
Chaitophorus nigrae
S. kinuyanagi Cavariella nipponica
Chaitophorus horii, saliapterus
S. koreensis Cavariella salicicola
Tuberolachnus salignus
S. koriyanagi Aphis farinosa
Cavariella aegopodii, konoi,
nipponica
Chaitophorus saliapterus,
salijaponicus
Plocamaphis coreana
[Pterocomma salijaponica]
Tuberolachnus salignus
S. laevigata Cavariella pastinacae
Chaitophorus macrostachyae,
452 Salix

monelli, nigrae, salicorticis

Fullawaya saliciradicis
Macrosiphum californicum
Tuberolachnus salignus
S. lanata Cavariella archangelicae
Chaitophorus truncatus
Phylloxerina ?capreae
S. lapponum [Acyrthosiphon aurlandicum]
Aphis farinosa
Cavariella aquatica
Chaitophorus lapponum
Plocamaphis amerinae ssp.
borealis
Pterocomma rufipes, salicis
S. lasiandra Cavariella konoi, pastinacae
(Pacific Willow) Macrosiphum californicum
Pterocomma ?populeum, smithiae
Tuberolachnus salignus
S. lasiolepis Cavariella aegopodii
(Arroyo Willow) Chaitophorus monelli, nigrae
Macrosiphum californicum
Phylloxerina salicicola
Plocamaphis flocculosa
Tuberolachnus salignus
S. laurina Chaitophorus niger
S. livida Chaitophorus niger, truncatus
S. longifolia Cavariella aegopodii
Chaitophorus abditus, nigrae,
viminalis
Fullawaya saliciradicis
Pterocomma konoi, salicis
S, longipes = S. caroliniana
S. lucida Cavariella salicis
(Shining Willow) Chaitophorus viminalis
Pterocomma salicis
S. lutea Aphis farinosa
Macrosiphum californicum
Pterocomma bicolor
S. macrostachya Chaitophorus crucis,
macrostachyae
Tuberolachnus salignus
 Salix 453

S. matsudana Stomaphis sinisalicis

S. medenii Cavariella aegopodii, archangelicae
S. melanopsis Chaitophorus macrostachyae
S. miyabeana Cavariella salicicola
S. multinervis Chaitophorus salijaponica
S. myrsinifolia Chaitophorus lapponum
S. nigra Aphis farinosa
(Black Willow) Cavariella aegopodii, salicis
Chaitophorus nigrae, viminalis,
viminicola
Pterocomma smithiae
S. nigricans Aphis farinosa
Cavariella aquatica, archangelicae
Chaitophorus lapponum,
nigricantis, vitellinae
Plocamaphis flocculosa ssp.
goernitzi
Pterocomma populeum, salicis
S. parallelinervis Plocamaphis coreana
S. pentandra Cavariella aegopodii,
(Bay Willow, Sweet Willow) archangelicae, pastinacae,
salicicola
Chaitophorus niger, pentandrinus,
truncatus, vitellinae
Pterocomma konoi, pilosum,
rufipes, salicis
S. phylicifolia Aphis farinosa
Cavariella intermedia, konoi
Chaitophorus lapponum,
nigricantis, salijaponicus ssp.
stroyani, vitellinae
Pterocomma rufipes
S. polaris Pemphigus saliciradicis
S. pseudolasiogyne Cavariella salicicola
Chaitophorus saliciniger
Tuberolachnus salignus
S. purpurea Aphis farinosa, roumanica
(Purple Osier) Cavariella aegopodii, aquatica,
archangelicae, konoi, rutila,
salicicola
454 Salix

Chaitophorus beuthani,
diversisetosus ssp. austriacus,
mordvilkoi, niger, salicti,
truncatus, vitellinae
Plocamaphis amerinae
Pterocomma italica, konoi,
pilosum, salicis
Tuberolachnus salignus
S. pyrenaica Aphis farinosa
S. repens Aphis farinosa
Cavariella aegopodii
Chaitophorus hypogaeus, niger,
parvus, salicti
Neopterocomma verhoeveni
Plocamaphis flocculosa ssp.
goernitzi
Pterocomma jacksoni, konoi,
pilosum
S. reticulata [Pemphigus groenlandicus]
S. rorida Aphis farinosa ssp. yanagicola
Cavariella konoi, nipponica,
salicicola
Chaitophorus horii, saliapterus
S. russica Chaitophorus brunealineatus
S. sachalinensis Aspidophorodon salicis
Cavariella nipponica
Chaitophorus horii
Prociphilus oriens
S. salicifolium Pterocomma tuberculatum
S. schwerinii Cavariella nipponica
Chaitophorus horii, niger
Pterocomma konoi, rufipes
S. scouleriana Aphis farinosa
Macrosiphum californicum
Pterocomma salicis
S. sieboldiana Cavariella nipponica
Chaitophorus matsumurai
S. silesiaca Pachypappa warschavensis
S. sitchensis Aphis farinosa
(Sitka Willow) Pterocomma smithiae
Tuberolachnus salignus
 Salix 455

S. smithiana (incl. var. velutina) Chaitophorus beuthani

Plocamaphis flocculosa ssp.
macrosiphon
Pterocomma konoi
Tuberolachnus salignus
S. subfragilis Cavariella salicicola
S. tetrasperma Cavariella aegopodii
(Indian Willow, Jalmola) Chaitophorus himalayensis,
pakistanicus
Lachnus salicis
Tuberolachnus salignus
S. tianschanica Chaitophorus salicti
Pterocomma salicis
S. triandra Aphis farinosa
(Almond-Leaved Willow) Cavariella aegopodii, konoi,
salicicola
Chaitophorus beuthani, niger,
nigricantis, truncatus, vitellinae
Macrosiphum californicum
Plocamaphis flocculosa
Pterocomma jacksoni, konoi,
pilosum, rufipes, salicis
Tuberolachnus salignus
S, udensis Cavariella japonica, nipponica,
salicicola
Chaitophorus horii, saliapterus
Elatobium hidaense
Pterocomma konoi, rufipes
S. urbaniana Cavariella aquatica
[Pemphigus yanagi Shinji, 1928]
S. viminalis Aphis farinosa
(Basket Willow, Common Osier) Cavariella aegopodii
Chaitophorus beuthani, capreae,
horii, niger, viminalis, vitellinae
Plocamaphis amerinae
Pterocomma konoi, pilosum,
rufipes, salicis
Stomaphis longirostris
Tuberolachnus salignus
S. vitellina Cavariella aegopodii, theobaldi
(Yellow Willow) Chaitophorus gomesi, niger,
salicti, truncatus, vitellinae
Pterocomma pilosum, salicis
456 Salix

S. wallichiana Prociphilus (Neoparacletus) ghanii

S. warburgii Aphis gossypii
Cavariella araliae, salicicola
Toxoptera aurantii
Tuberolachnus salignus
S. wardiana Cavariella archangelicae
S. xerophila [Pterocomma xerophilae
Ivanoskaya, 1971]
S. yezoalpina Tuberolachnus salignus
Salix spp. (unidentified) Aleurodaphis sinisalicis
Aspidophorodon harvensis,
sinisalicis
[Bituberculaphis inexpectata
Rusanova, 1943]
Cavariella aspidaphoides, biswasi,
digitata, [hidaensis], japonica,
pustula, takahashii
Chaitophorus crinitus, eoessigi,
hokkaidensis, israeliticus,
knowltoni, manaliensis,
nigricentrus, nigritus,
pustulatus, remaudierei,
[salicifoliae] saliciniger,
shaposhnikovi, [tumurensis
Zhang in Zhang and Zhong,
1985a]
[Eriosoma yangi, yangiparasitica]
Fullawaya bradleyi, braggii,
bulbosa, terricola
Lachnus tatakaensis, yunlangensis
Longistigma xizangensis
Paducia antennata, aterrima
Plocamaphis [assetacea], martini
Pterocomma baicalense,
groenlandica, [henanense],
[salicicola Uhler, 1862],
salicis ssp. rohdendorfi,
[tibetasalicis]
[Tranaphis syriaca Börner ex
Bodenheimer, 1937]

Main key to willow-feeding aphids (apterous viviparae except where otherwise

stated)
1 Body very small (BL less than 1.24 mm), pear-shaped, broadest anteriorly.
Antenna 3-segmented. Spinal, pleural and marginal wax pore plates on
all thoracic and abdominal segments. (All morphs oviparous) . . . . . . 2
 Salix 457

Fig. 100. Head and prothorax, with detail of structure of wax pore plates, of aptera of A, Phylloxerina
salicis and B, Ph. capreae.

- BL usually more than 1.25 mm. Antenna 5- or 6-segmented. Wax glands

present or absent but never in a complete series on all segments. (Parth-
enogenetic females viviparous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 R IV+V elongate, 5-6 x its basal width and a little longer than antenna.
Facets of wax pore plates polygonal with subdivisions (Fig. 100A). Pro-
thorax usually with a hair-bearing supplementary wax pore plate between
the spinal and pleural wax pore plates on each side, sometimes fused with
the spinal wax pore plate, which then appears to be extended laterally and
includes 2 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phylloxerina salicis
- R IV+V 2.5-3.5 x its basal width and a little shorter than antenna. Facets
of wax pore plates more rounded (Fig. 100B). Prothorax usually without
a supplementary wax pore plate between the spinal and pleural wax pore
plates . . . . . . . . . . . . . Phylloxerina capreae or daphnoidis or salicicola*
3 Tergum sclerotic, warty, with a continuous marginal row of tubercles/wax
glands. Anal plate bilobed. R IV+V more than 2 x ANT III . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleurodaphis sinisalicis
- Tergum membranous or sclerotic but without a continuous marginal row
of tubercles. Anal plate entire. R IV+V less than 2 x ANT III . . . . 4
4 ANT PT/BASE less than 0.7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- ANT PT/BASE more than 0.7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
5 SIPH present as large pores on broad pigmented cones with numerous
hairs. Eyes multifaceted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
458 Salix

- SIPH absent. Eyes as triommatidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

6 Rostrum much longer than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Rostrum much shorter than body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 ANT VI a little shorter than ANT V. ANT III with 1-12 secondary
rhinaria, and ANT IV with 4-10 . . . . . . . . . . . . . Stomaphis longirostris
- ANT VI about 1.6 x ANT V. ANT I I I without secondary rhinaria and
ANT IV with 1-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . Stomaphis sinisalicis*
8 Abdomen with a large dark spinal tubercle . . . . Tuberolachnus salignus
- Abdomen without a spinal tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 BL more than 5.5 mm. Alata with pterostigma elongate, curved around
tip of forewing . . . . . . . . . . . . . . . . . . . . . . . . . . .Longistigma xizangensis*
- BL less than 5.5 mm. Alata (where known) with short pterostigma . . . 10
10 Dorsal abdomen with transverse rows of large black circular sclerites,
about 6 per tergite . . . . . . . . . . . . . . . . . . . . . . . . . . . Lachnus tatakaensis
- Dorsal abdomen without transverse rows of round black sclerites ... 11
11 ANT III (of aptera) with 11-15 secondary rhinaria, IV with 4-8 and V
with 0-2. R IV with 24-32 accessory hairs . . . Lachnus yunlangensis*
- ANT III with 0-1 secondary rhinaria, IV with 0-4 and V with 0-1. R IV
with about 10 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 R IV+V 0.42-0.51 x HT II . . . . . . . . . . . . . . . . . . . . . . . Lachnus salicis
- R IV+V 0.61-0.68 x HT II . . . . . . . . . . . . . . . . . Lachnus longirostrum
13 Hairs on antennae and legs long; e.g. those on ANT I I I longer than basal
diameter of segment. Abdomen with only marginal wax pore plates well
developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- Hairs on appendages short and sparse. Posterior abdominal tergites with
well-developed spinal and pleural wax pore plates . . . . . . . . . . . . . . . 15
14 Body hairs short, hairs on antennae and legs very long and fine. R IV+V
about as long as HT II. Marginal abdominal wax pore plates well devel-
oped. Anal plate forming a ring around CAUDA . . . Prociphilus oriens
- Hairs on body, legs and antennae of similar size, all with rather thick
bases. R IV+V 1.4-1.5 X HT II. Marginal abdominal wax pore plates
poorly developed, possibly represented by small round membranous areas
on each tergite. Anal plate wholly below CAUDA . . . Prociphilus ghanii
15 R IV+V acute, longer than HT II, which is without spinules and appears
fused to HT I. Antenna 5-segmented, with PT shorter than its basal
diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pachypappa warschavensis
- R IV+V blunt, shorter than HT II which is spinulose and has a functional
articulation with HT I. Antenna 6-segmented, with PT at least as long as
its basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pemphigus saliciradicis
16 Body usually rather densely clothed with either long and fine, or thick,
 Salix 459

Fig. 101. A, SIPH of Aspidophorodon harvensis; front of head of B, A. salicis and C, A. harvensis.

hairs; if hairs are rather short and sparse mid-dorsally then the antennae,
legs and CAUDA are all densely hairy . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Body hairs sparse and usually rather short, at least dorsally . . . . . . 18
17 Small to medium-sized (BL 1.5-2.6 mm). SIPH stump-shaped, usually with
reticulate sculpturing (see Fig. 103). CAUDA either knobbed, rounded or
bluntly triangular, with 6-14 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaitophorus spp. - KEY A (p. 460)
- Medium to large (BL 2.2-4.5 mm). SIPH almost cylindrical or swollen,
usually without reticulate sculpturing (see Fig. 106). CAUDA either
rounded or bluntly triangular with about 20-60 hairs, or tongue-shaped
with a constriction and fewer hairs (in which case SIPH are dark and
strongly swollen and antennae are 4-segmented) . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocommatini spp. - KEY B (p. 467)
18 ABD TERG 8 with a backwardly-directed supracaudal process bearing a
pair of hairs near apex (Fig. 107) . . . Cavariella spp. - KEY C (p. 472)
- ABD TERG 8 without a supracaudal process . . . . . . . . . . . . . . . . . . . 19
19 Antennal tubercles with forwardly-directed processes on either side of a
well-developed median tubercle, and ANT I also with projecting inner
face. Antenna 4- or 5-segmented. SIPH thin and flangeless, spoon-
shaped, obliquely truncated at apex (Fig. 101A) . . . . . . . . . . . . . . . . . 20
- Antennal tubercles developed or undeveloped, without processes. Antenna
usually 6-segmented. SIPH not spoon-shaped, with a flange . . . . . . . 21
20 Processes on antennal tubercles not much longer than their basal widths,
only slightly longer than median tubercle, which is bilobed (Fig. 101B).
Hairs on frontal processes, and on ABD TERG 8, blunt or pointed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aspidophorodon salicis
- Processes on antennal tubercles more than 2 x longer than median
tubercle, which is not bilobed (Fig. 101C). Hairs on frontal processes and
ABD TERG 8 slightly capitate . . . . . . . . . . . Aspidophorodon harvensis
21 SIPH usually dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
460 Salix

- SIPH pale at least at base, if rather dark then with subapical polygonal
reticulation ..................................................23
22 ANT PT/BASE 3.5 or more. CAUDA dark with 10-26 hairs. Stridulatory
apparatus present (Fig. 122) . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- ANT PT/BASE 2.0-3.2. CAUDA paler than SIPH, with 4-9 hairs. No
Stridulatory apparatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
23 Antennal tubercles undeveloped. ANT PT/BASE 1.5-2.3. CAUDA dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
- Antennal tubercles moderately to well developed, broadly divergent (e.g.
Fig. 42K). ANT PT/BASE 2.1-7.5. CAUDA pale . . . . . . . . . . . . . . . . 25
24 R IV+V 1.03-1.25 x HT II. ABD TERG 2-6 each with 2-3 marginal hairs
on each side. ABD TERG 8 with 2, or rarely 3-4, hairs . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis farinosa
- R IV+V 0.93-1.07 x HT II. ABD TERG 2-6 each with 5-11 marginal hairs
on each side. ABD TERG 8 with 4-7 hairs . . . . . . . . . Aphis roumanica
25 ANT PT/BASE 6.5-7.5. SIPH darkened distally with a subapical zone of
polygonal reticulation . . . . . . . . . . . . . . . . . . . Macrosiphum californicum
- ANT PT/BASE 2.1-4.5. SIPH wholly pale with no distinct zone of
polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 ANT III with 1-5 secondary rhinaria. ANT PT/BASE 3.5-4.5. SIPH
1.00-1.32 x CAUDA . . . . . . . . . . . . . . . . . Acyrthosiphon aurlandicum*
- ANT III without rhinaria. ANT PT/BASE 2.1-2.6. SIPH 2.5-3.5 x
CAUDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27 Antenna more than 0.75 x BE, with ANT III longer than head width
across (and including) eyes . . . . . . . . . . . . . . . Neoacyrthosiphon dubium
- Antenna less than 0.7 x BL, with ANT III about 0.5 x head width across
eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elatobium hidaense

KEY A - Chaitophorus species on Salix (apterous viviparae, except where

otherwise stated)
1 CAUDA broadly rounded, less than 0.6 x its basal width, without a trace
of a constriction (e.g. Fig. 102A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- CAUDA either with a distinctly knobbed apex, or bluntly conical with or
without a slight mid-way constriction; more than 0.6 x its basal width in
dorsal view (e.g. Figs 102B-D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2 ANT BASE VI with 7-14 hairs. R IV with 11-20 accessory hairs. ANT
PT/BASE usually less than 1 . . . . . . . . . . . . . . . . . macrostachyae group
- ANT BASE VI with 2-5 hairs. R IV with 2-8 accessory hairs. ANT
PT/BASE usually more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
 Salix 461

Fig. 102. CAUDA of aptera of A, Chaitophorus saliciniger, B, Ch. viminalis, C, Ch. nigrae, D, Ch. vitellinae;
ANT III of aptera of E, Ch. crucis, F, Ch. saliciniger and G, Ch. capreae.

3 Tergum ornamented with numerous small rounded nodules . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pustulatus
- Tergum with mainly imbricate and at least partly reticulate ornamentation
............................................................. 4
4 Hind femora much darker than tergum. R IV with 6-8 accessory hairs.
Dorsal pigmentation rather uniform with darker intersegmental markings
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . utahensis
- Hind femora as pale as or paler than tergum. R IV with 2-4 accessory hairs.
Dorsal pigmentation either pale or only regionally pigmented . . . . . . . . 5
5 Tergum entirely pale. Hairs on ANT III mainly thick, stiff and blunt
(Fig. 102E). First tarsal segments with 5 hairs . . . . . . . . . . . . . . . . crucis
- Tergum regionally pigmented. Hairs on ANT III mostly long and fine-
pointed (e.g. Fig. 102F). First tarsal segments with 7 (or rarely 6) hairs
............................................................. 6
6 Tergum dark laterally and usually with a distinct pale central area . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . saliciniger
- Tergum paler laterally, with dark brown central area . . . . nigricentris
7 ANT III with only 0-4 hairs, the longest of which are only 5-20 µm long
(Fig.102G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- ANT III with 2-30 hairs, but if 4 or less then the longest are 30-100 µm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
462 Salix

Fig. 103. Hairs on ABD TERG 4-6 of aptera of A, Chaitophorus capreae and B, Ch. horii; C, basal part
of hind tibia of apterous vivipara of Ch. saliniger; ABD TERG 5 and 6 of aptera of D, Ch. nigrae and
E, Ch. pentandrinus.

8 BL 1.8-2.1 x maximum body width. ABD TERG 1-5 each with only 0-12
smaller accessory hairs between the long spinal, pleural and marginal
pairs, and mostly forming a single row with them (Fig. 103A) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . capreae
- Body narrower; BL 2.1-2.5 x maximum body width. ABD TERG 1-5
each with 12-30 smaller accessory hairs, some in an additional row
anterior to the large primary hairs (Fig. 103B) . . . . . . . . . . . . . . . . horii
9 Abdominal tergum membranous and mainly pale except for dusky
scleroites at bases of spinal and pleural hairs, lateral sclerites and trans-
verse bands on ABD TERG 7 and 8. Dorsal body hairs very long and
pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hypogeus
- Abdominal tergum sclerotic, pale or pigmented, with usually at least ABD
TERG 2-6 fused together. Dorsal body hairs variable . . . . . . . . . . . 10
10 Hind tibia slightly swollen ventrally near base, with a group of 2-17
small circular ?scent plaques (Fig. 103C). Tergum dark, with nodulose
ornamentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . saliniger
 Salix 463

- Hind tibia without a basal swollen part bearing ?scent plaques. Tergum
pale or dark, ornamentation variable . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 CAUDA bluntly conical, often with an indentation or constriction
partially delimiting a tongue-shaped or rounded apical part, but with the
width at the constriction not usually less than 0.8 x maximum diameter
of apical part (e.g. Figs 102C, D; examine several specimens) . . . . . 12
- CAUDA always with a distinct constriction or heck dividing it into a
triangular basal part and a globular apical knob, with the width at the
constriction less than 0.8 x maximal diameter of the knob (Fig. 102B)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
12 ANT BASE VI with 2-4 hairs, but usually 3 hairs (examine several
specimens). Tergum ornamented with separate small nodules. (R IV+V
1.05-1.20 x HT II; ANT PT/BASE 1.5-2.2; dorsal body hairs long,
100-160 µm; dorsum pale in spring and early summer, with dark pleural
bands in late summer) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vitellinae
- ANT BASE VI with 2 hairs, or rarely 3 hairs, and without the above com-
bination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Tergum pale. Dorsal body hairs mostly with furcate apices . . . . . . . 14
- Tergum usually dark, either t o t a l l y or with a paler spinal region. Dorsal
body hairs usually with pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . 16
14 R IV+V less than 0.8 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . monelli
- R IV+V at least 0.9 x HT I I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 ANT PT/BASE 1.0-1.3, R IV+V 1.1-1.3 x HT II . . . . shaposhnikovi
- ANT PT/BASE 2.1-2.5, R IV+V 0.9-1.1 x HT II . . . . . . . . furcatus
16 ABD TERG 1-6 all fused . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- ABD TERG 1 separate, 2-6 fused or separate . . . . . . . . . . . . . . . . . . 18
17 SIPH unpigmented, with a clear surrounding area. ABD TERG 1-8 all
with pale spinal areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pallipes
- SIPH dusky, without any clear surrounding area (Fig. 103D). ABD TERG
1-5 often with clear spinal areas in heavily pigmented specimens, but then
ABD TERG 7 and 8 are always evenly pigmented . . . . . . nigrae group
18 ABD TERG 2 separate from 3, and ABD TERG 3-6 often with separate
or only partially fused bands. R IV+V 1.0-1.3 x HT II . . . . . ramicola
- ABD TERG 2-6 usually wholly or partially fused; if separate in some
specimens then R IV+V 0.8-1.0 x HT II . . . . . . . . . . . . . . . . . . . . . . 19
19 R IV with 3-6 accessory hairs (most commonly 4). First tarsal segments
most commonly with 5 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . parvus
- R IV with 2 accessory hairs. First tarsal segments most commonly with
7 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . mordvilkoi
464 Salix

20 ABD TERG 1 fused with 2-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

- ABD TERG 1 separate from 2-6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
21 ABD TERG 1-6 with a pattern of reticulate imbrication, at least on
pleural areas. Dorsal abdominal hairs mostly long and pointed (Fig. 103D)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nigrae group
- Ornamentation of ABD TERG 1-6 not reticulate, usually consisting of
separate small nodules or spicules, sometimes smooth. Dorsal abdominal
hairs pointed or bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 R IV+V short, less than 1.5 x its basal width, 0.6-0.8 x HT II . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
- R IV+V more than 1.5 x its basal width, 0.9-1.4 x HT II . . . . . . 24
23 First tarsal segments with usually 5 hairs, rarely 6 on some tarsi. Body
and appendages always pale. Hairs on ANT VI BASE rather short, the
longer of the two up to about 2 x basal diameter of ANT III . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pusillus
- First tarsal segments with 6-7 hairs (rarely 5 on some tarsi). Body and
appendages variably pigmented. Longest hair on ANT BASE VI more
than 2.5 x basal diameter of ANT III . . . . . . . . . . . . . . . . . . . viminalis
24 Antennae usually 5-segmented, with ANT BASE V usually bearing only
a single, short hair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . minutus
- Antennae 6-segmented, ANT BASE VI with 2-3 hairs . . . . . . . . . . . 25
25 R IV+V 0.90-0.95 x HT II, and with only 2 accessory hairs. Dorsal
abdominal hairs mostly with bifurcate apices . . . . . . . . . . . . . . . eoessigi
- R IV+V 1.0-1.4 x HT II, with 2-15 accessory hairs. Dorsal abdominal
hairs with either bifurcate or pointed apices . . . . . . . . . . . . . . . . . . . . 25
26 All femora dark. R IV+V 1.2-1.4 x HT II, with 3-15 accessory hairs.
Dorsal abdominal hairs mostly with bifurcate apices . . . . . . . . longipes
- Either all femora pale, or fore femora paler than mid- and hind femora.
R IV+V 1.0-1.2 x HT II, with 2-4 accessory hairs. Dorsal abdominal
hairs mostly with pointed apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
27 Mid- and hind femora very dark. ANT PT/BASE 3.0-4.2 . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viminicola
- Mid- and hind femora pale or dusky. ANT PT/BASE 2.2-2.8 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hokkaidensis
28 First tarsal segments usually with 6-7 hairs (but ocasionally 5 on some or
even all tarsi of some specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
- First tarsal segments normally with 5, occasionally 6, hairs . . . . . . . 38
29 ANT VI BASE with 3-4 hairs which have blunt or expanded apices.
 Salix 465

R IV+V 1.05-1.20 x HT II, with R IV bearing 5-7 accessory hairs ...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vitellinae ssp. danubicus
- ANT VI BASE with 2-3 hairs with pointed apices. R IV+V 0.60-1.05 x
HT II, with R IV bearing 2-6 accessory hairs . . . . . . . . . . . . . . . . . . . 30
30 SIPH dark, at least apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
- SIPH pale, even when tergum is dark . . . . . . . . . . . . . . . . . . . . . . . . . . 33
31 R IV with 4-6 accessory hairs. ANT PT/BASE 3.0-3.8 . . . . . . . . . . . . .
................................................... himalayensis
- R IV with 2-3 accessory hairs. ANT PT/BASE 1.8-3.3 . . . . . . . . . . 32
32 Spinal hairs with blunt or expanded apices, shorter than marginal hairs
on same tergites. SIPH reticulated on distal half only. Tergum rather pale
or with dark brown pleural longitudinal stripes . . . . . . . . diversisetosus
- Spinal hairs long and pointed and similar in length to marginal hairs.
SIPH almost wholly reticulated. Tergum with transverse bars, usually
fused between ABD TERG 3-6 . . . . . . . . . . . . . . . . . . . . . . remaudierei*
33 Longest hair on ANT BASE VI (or V if antenna 5-segmented) more than
1.5 x maximum diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . 34
- Longest hair on ANT BASE VI less than 1.5 x maximum diameter of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
34 ANT PT/BASE 3.4-3.8. All dorsal abdominal hairs long and fine-
pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . manaliensis
- ANT PT/BASE 2.1-3.0. Spinopleural abdominal hairs thick with
abruptly pointed, blunt or bifid apices (Fig. 103E), only marginal hairs
sometimes long and fine-pointed . . . . . . . . . . . . . . . . . . . . . pentandrinus
35 ABD TERG 1-5 with longest spinopleural hairs 2-4 x basal diameter of
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
- ABD TERG 1-5 with longest spinopleural hairs 4-8 x basal diameter of
ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
36 ANT V with 1-3 hairs. R IV with 2 accessory hairs. (Alata with secondary
rhinaria on ANT III-V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . saliapterus
- ANT V with 3-6 hairs. R IV with 2-4 accessory hairs. (Alata with secon-
dary rhinaria only on ANT III) . . . . . . . . . . . . . . . . . quinquemaculatus
37 Antennae wholly pale except for duskiness around primary rhinaria.
(Alata with broad transverse dark bars on ABD TERG 3-8, often fused
into a solid patch on ABD TERG 3-6) . . . . . . . . . . . . . . . . . nigricantis
- ANT I and most of ANT V and VI darker than ANT III. (Alata with
separate, narrow, often broken transverse bars on ABD TERG 3-8) .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . truncatus
466 Salix

Fig. 104. SIPH and surrounding cuticle of aptera of A, Chaitophorus salijaponicus and B, Ch. niger.

38 R IV with 3-6 (usually 4) accessory hairs . . . . . . . . . . . . . . . . . . . . . . . 39

- R IV with 2, or rarely 3-4, accessory hairs . . . . . . . . . . . . . . . . . . . . . 40
39 R IV+V 0.7-1.0 x HT I I . Legs, or at least the femora, pigmented
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lapponum
- R IV+V 1.1-1.6 x HT II. Legs except tarsi wholly pale . . . . . . salicti
40 Antennae and legs except tarsi wholly pale, contrasting with blackish-
brown tergum; at most, ANT VI BASE and hind femora slightly dusky
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nigritus
- Tergum pale or dark; if very dark, then at least ANT VI BASE and hind
femora are also dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
41 Dorsal abdominal hairs all fine-pointed . . . . . . . . . . . . . . . . . . . . . . . . 42
- Dorsal abdominal hairs mostly with either abruptly acute, blunt, expanded
or bifurcate apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
42 Tergum pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . israeliticus
- Tergum dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
43 ANT V with only 1 hair in most specimens. SIPH completely or partially
included in dorsal shield, without a clear ring at base (Fig. 104A) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . salijaponicus
- ANT V with 1-4 hairs. SIPH usually with a broad, clear basal ring
separating them from dorsal shield (Fig. 104B) . . . . . . . . . . . . . . . . niger
44 Dorsal abdominal hairs mostly with abruptly acute, blunt, or narrowly or
minutely bifid apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
- Dorsal abdominal hairs mostly with distinctly expanded, bifurcate or fan-
shaped apices, often as broad apically as at base . . . . . . . . . . . . . . . . 47
 Salix 467

45 Tergum dark. ANT PT/BASE 2.5-3.3 . . . . . . . . . . . . . . . . . . . . stroyani

- Tergum pale. ANT PT/BASE 1.2-2.8 . . . . . . . . . . . . . . . . . . . . . . . . . 46
46 R IV+V 0.11-0.13 mm, 1.1-1.4 x HT II . . . . . . . . . . . . . . . . . . gomesi
- R IV+V 0.09-0.11 mm, 0.8-1.0 x HT II . . . . . . . . . . . . . pakistanicus
47 ANT PT/BASE 1.3-2.0; R IV+V 1.1-1.5 x HT II . . . . . . . . . . . . . 48
- ANT PT/BASE 2.6-4.0; R IV+V 0.8-1.2 x HT II . . . . . . . . . . . . . 49
48 Antennae 6-segmented. ANT V usually with 1 hair, and VI BASE with
1-2 short hairs, the longest not or hardly exceeding basal diameter of ANT
III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crinitus*
- Antennae usually 5-segmented; penultimate segment (IV or V) with 1-4
hairs, and ANT V (or VI) BASE with 2 hairs, the longest 1.7-2.5 x basal
diameter of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . similis*
49 ANT PT/BASE 3.5-4.0; R IV+V 0.8-0.9 x HT II . . . . . longiunguis
- ANT PT/BASE 2.6-3.0; R IV+V 1.05-1.11 x HT II . . . . matsumurai

KEY B - Pterocommatini species on Salix (can be applied to both apterous

and alate viviparous females)
1 Antennae 4-segmented. SIPH blackish, large and strongly inflated, with
distal imbrication (Fig. 106A). CAUDA clearly longer than its basal width,
with a midlength constriction, and bearing 5-14 hairs . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paducia antennata
- Antennae 6-segmented. SIPH pale or dark, rather small, variously shaped,
sometimes absent. CAUDA shorter or longer than its basal width, rounded
or bluntly triangular, with 20-60 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH present or absent, when present rounded at apex and completely
lacking a flange (e.g. Figs 106B-F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- SIPH truncate apically, usually with at least a small flange (e.g.
Figs106I-P) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
3 SIPH very short, conical with a broad base (e.g. Fig. 106B). Dorsal hairs
rather sparse and mostly shorter than basal diameter of ANT III .... 4
- SIPH either longer their basal width, with a constricted base (e.g.
Fig. 106C), vestigial or absent. Dorsal hairs all longer than basal diameter
of ANT III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4 ANT PT/BASE more than 1. R IV with 16-20 accessory hairs. CAUDA
a little shorter than its basal width . . . . . . . . Neopterocomma asiphum
- ANT PT/BASE less than 1. R IV with 7-10 accessory hairs. CAUDA a
little longer than its basal width . . . . . . . . Neopterocomma verhoeveni
5 Empodial hairs short, less than half of length of claws (Fig. 105A).
468 Salix

Fig. 105. Empodial hairs of A, Plocamaphis flocculosa and B, Fullawaya saliciradicis; prothoracic
marginal tubercle (right side) of aptera of C, Fullawaya bulbosa (after Richards, 1966b), and
D, Pterocomma bicolor; E, ANT VI of Pterocomma chaetosiphon.

Mid-dorsal abdominal hairs rather sparse, mostly shorter than the

distances between their bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Empodial hairs long, much more than half of length of claws (Fig. 105B).
Mid-dorsal abdominal hairs dense and longer than the distances between
their bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
6 SIPH up to about 0.5 x HT II (e.g. Fig. 106C). Dark markings only on
ABD TERG 1 and 2, or absent. Abdominal marginal tubercles present or
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- SIPH about as long as HT II or longer (e.g. Figs 106E, F). Abdomen with
paired dark patches on ABD TERG 2-5. Abdominal marginal tubercles
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7 Hairs on ANT III mostly shorter than basal diameter of segment. R IV+V
shorter than SIPH, and bearing only 2-3 accessory hairs . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis coreana
- Hairs on ANT III mostly much longer than basal diameter of segment.
R IV+V longer than SIPH and bearing 4-6 accessory hairs . . . . . . . . 8
8 Abdomen with small marginal tubercles on most tergites . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis amerinae
- Abdomen usually without any marginal tubercles . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis martini
9 SIPH less than 1.25 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
 Salix 469

- SIPH more than l .25 X HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

10 SIPH (Fig. 106E) usually more than 1.1 X HT II. Hairs on ANT III
mostly about 1.5 x basal diameter of segment or shorter, rarely a few
hairs up to 2 or more x basal diameter . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis flocculosa flocculosa
- SIPH (Fig. 106F) less than 1.1 x HT II. Hairs on ANT III mostly more
than 2 x basal diameter of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis flocculosa brachysiphon
11 SIPH 1.25-1.5 x HT II. ANT III with 12-14 secondary rhinaria . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis flocculosa goernitzi
- SIPH more than 1.5 x HT II. ANT III with 15-18 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plocamaphis flocculosa macrosiphon
12 Marginal tubercles on prothorax bulbous, with a narrow neck (Fig. 105C).
Abdominal marginal tubercles either similar or tubercular with height
greater than basal diameter . . . . . . . . . . . . . . . . . . . . Fullawaya bulbosa*
- Prothoracic and abdominal marginal tubercles broad and low . . . . 13
13 SIPH absent, or present only as vestigial, misshapen tubercles without
apical pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
- SIPH present, complete with an apical pore, even if very small (e.g.
Figs106G, H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14 Hairs on ANT III maximally 1.5 x basal diameter of segment . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fullawaya saliciradicis
- Hairs on ANT III maximally 2 or more x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fullawaya bradleyi (part)
15 SIPH (Fig. 106H) very small, less than 0.5 X HT II Fullawaya braggii
- SIPH more than 0.5 x HT II (e.g. Fig. 106G) . . . . . . . . . . . . . . . . . . 16
16 Rostrum long, extending beyond 5th abdominal segment; R IV with 12-13
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fullawaya terricola
- Rostrum not reaching 5th abdominal segment; R IV with 15-23 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fullawaya bradleyi (part)
17 SIPH cylindrical or slightly tapering, bearing 1-4 hairs on basal half
(Fig. 106I). ANT PT/BASE less than 1.5, with PT very thick, only a little
narrower than BASE VI, and bearing many long hairs (Fig. 105E) .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma chaetosiphon
- SIPH swollen or cylindrical, without hairs. ANT PT/BASE more or less
than 1.5, with PT much narrower than BASE and bearing a few short
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
470 Salix

Fig. 106. SIPH of aptera of A, Paducia antennata, B, Neopterocomma verhoevenei, C, Plocamaphis

coreana, D, P. martini, E, P. flocculosa, F, P. flocculosa brachysiphon, G, Fullawaya terricola,
H, F. braggi, I, Pterocomma chaetosiphon, J, Pt. salicis, K, Pt. jacksoni, L, Pt. italica,
M, Pt. sanguiceps, N, Pt. pilosum, O, Pt. bicolor, P, Pt. rufipes (all to same scale).
 Salix 471

18 ANT II with 8-20 hairs. ANT III almost always with a few (1-13) second-
ary rhinaria. SIPH slightly to markedly swollen . . . . . . . . . . . . . . . . . 19
- ANT II with 3-6 hairs. ANT III never with secondary rhinaria. SIPH
swollen or cylindrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
19 ANT PT/BASE 0.7-1.0; R IV+V 0.65-0.90 X HT II. SIPH strongly
swollen, to more than 1.5 x their basal width (Fig. 106J) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma salicis
- ANT PT/BASE 0.8-1.9 (only below 1.0 in spring); R IV+V 0.85-1.30 X
HT II. SIPH slightly to moderately swollen, up to 1.5 x basal width
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 SIPH moderately swollen, with maximal swelling distal to midlength, and
with imbrication reduced on swollen part, the imbrication throughout
showing no trace of spinules (Fig. 106K). ANT PT/BASE 0.8-1.3 in
spring and 1.25-1.65 in summer/autumn . . . . . . Pterocomma jacksoni
- SIPH slightly swollen, with maximal swelling at or proximal to midlength,
and with rather evenly distributed spinulose imbrication (Fig. 106L). ANT
PT/BASE 1.2-1.8 in spring and 1.5-1.9 in summer/autumn . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma italica
21 SIPH short and stout, with maximal swelling near base (Fig. 84A). ANT
PT/BASE 1.0-1.2 . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma smithiae
- SIPH cylindrical or swollen distally. ANT PT/BASE 1.0-2.2 . . . . . 22
22 ANT BASE VI with 5-10 long hairs exceeding diameter of segment, plus
2-4 short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
- ANT BASE VI with only 1-3 long hairs, plus 2-4 short hairs . . . . 25
23 SIPH (Fig. 106M) a little shorter than HT II . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma sanguiceps
- SIPH (Fig. 106N) longer than HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Prothorax and ABD TERG 1-7 usually all with well-developed marginal
tubercles, rarely missing from some segments. ABD TERG 7 with 13-34
spinopleural hairs (i.e. not including marginal hairs near the marginal
tubercle and spiracle on each side). Alata with 22-38 secondary rhinaria
on ANT III, 0-3 on ANT IV . . . . . . . . . . Pterocomma pilosum konoi
- Prothorax and ABD TERG 1-7 without marginal tubercles, or rarely with
1-2 small ones. ABD TERG 7 with 10-20 spinopleural hairs. Alata with
33-44 secondary rhinaria on ANT III and 0-5 on IV . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma pilosum pilosum
25 Prothoracic and abdominal marginal tubercles small (not much larger
than a hair-base) or absent. ANT PT/BASE 1.4-1.8; SIPH 0.23-0.31 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma groenlandica
- Marginal tubercles present and well developed on prothorax and most of
ABD TERG 1-7; variably shaped but often conical and broad-based,
472 Salix

much larger than adjacent hair-bases (e.g. Fig. 105D). ANT PT/BASE
1.3-2.2; SIPH 0.22-0.45 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
26 Longest hairs on ANT III 85-115 µm, 2.2-3.3 x basal diameter of seg-
ment. SIPH usually markedly swollen distally, up to 1.5-1.9 x their basal
diameter (Fig. 106O). Alata with 12-26 secondary rhinaria on ANT III
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocomma bicolor
- Longest hairs on ANT III 120-150 µm, 3-4 x basal diameter of segment.
SIPH varying in shape, almost cylindrical or swollen to not more than 1.5
x their basal diameter (e.g. Fig. 106P). Alata with usually 25-30 secon-
dary rhinaria on ANT III . . . . . . . . . . . . . . . . . . . . . Pterocomma rufipes

KEY C - Apterae viviparae (except fundatrices) of Cavariella species on Salix

1 Tergum and SIPH blackish; antennae and legs also mainly dark . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nigra
- Tergum sclerotic but usually pale, if rather pigmented then antennae, legs
and SIPH are mainly pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH cylindrical or tapering, not at all swollen (e.g. Figs 108A, B) . . . 3
- SIPH clavate or at least slightly swollen (e.g. Figs 108C-J) . . . . . . . . 4
3 ANT PT/BASE 1.25-1.45. Process on ABD TERG 8 conical with 2
short hairs placed well below apex, plus 1-3 very small hairs more

Fig. 107. Supracaudal process of aptera (from Salix) of A, Cavariella digitata, B, C. theobaldi,
C, C. araliae, D, C. japonica, E, C. konoi, F, C. borealis, G, C. rutila (paratype), H, C. aegopodii,
I, C. aspidaphoides.
 Salix 473

0.1 mm

Fig. 108. SIPH (left side) of aptera (from Salix) of A, Cavariella digitata, B, C. theobaldi,
C, C. pastinacae, D, C. araliae, E, C. japonica, F, C. salicicola, G, C. konoi, H, C. aquatica, I, C. salicis,
J, C. aegopodii.

basad (Fig. 107A). SIPH strongly imbricated over entire length (Fig. 108A)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . digitata
- ANT PT/BASE more than 2.0. Process on ABD TERG 8 usually
quadrate, with 2 hairs at apical corners and none more basad (Fig. 107B).
SIPH more weakly imbricated, often almost smooth on basal half
(Fig.108B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . theobaldi
4 ANT PT/BASE more than 2.4. SIPH long, clavate, smooth or weakly
imbricated (Fig. 108C), process on ABD TERG 8 quadrate . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pastinacae
- ANT PT/BASE less than 2.4. SIPH various . . . . . . . . . . . . . . . . . . . . . 5
5 R IV+V more than 1.3 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- R IV+V less than 1.3 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
6 ANT PT/BASE less than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . nipponica
474 Salix

- ANT PT/BASE more than 1.2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7 Process on ABD TERG 8 short and broad, much shorter than CAUDA
(Fig. 107D). SIPH swollen over entire length, without a narrower central
section (Fig. 108E). Tergum often pigmented, ornamented with angular
pustules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . japonica
- Process on ABD TERG 8 long, with distal part finger-like, as long as or
longer than CAUDA. SIPH slightly clavate (i.e. with a narrow central
section; Fig. 108D). Tergum pale, ornamented with rounded pits . . . 8
8 Process on ABD TERG 8 very long and broad-based, with tip extending
beyond CAUDA (Fig. 107C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . araliae
- Process on ABD TERG 8 as long as or a little longer than CAUDA . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . biswasi
9 SIPH short and thick, swollen over most of length, less than 5 x their
maximum width (Fig. 108F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . salicicola
- SIPH slightly to moderately swollen distally, with narrower basal section;
more than 5 x longer than maximum width of swollen part (Figs 108G-J)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10 SIPH completely flangeless, rounded apically (Fig. 108H; absent from
fundatrices). CAUDA with 8-12 hairs . . . . . . . . . . . . . . . . . . . . aquatica
- SIPH with at least a small flange. CAUDA with 4-6 hairs . . . . . . . 11
11 ANT PT/BASE more t h a n 1.4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
- ANT PT/BASE less than 1.35 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
12 SIPH longer than head width across (and including) eyes. Process on ABD
TERG 8 at least 1.6 x maximum width of swollen part of SIPH. (Alata
with a complete black band on ABD TERG 6 and with 0-19 secondary
rhinaria on ANT IV) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . archangelicae
- SIPH equal to or a little shorter than head width across eyes (Fig. 108G).
Process on ABD TERG 8 (Fig. 107E) less than 1.5 x maximum width of
swollen part of SIPH. (Alata with a pair of roundish dark marks on ABD
TERG 6 and with 2-8 secondary rhinaria on ANT IV) . . . . . . . konoi
13 R IV with 2 accessory hairs in most specimens . . . . . . . . . . . . . . . . . . 14
- R IV without any accessory hairs in most specimens . . . . . . . . . . . . . 17
14 ANT PT/BASE less than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
- ANT PT/BASE more than 1.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
15 R IV+V more than 1.15 x HT II. (Alata with 14-32 secondary rhinaria
on ANT III and 2-6 on IV) . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedia
- R IV+V less than 1.15 x HT II. (Alata with 5-9 secondary rhinaria on
ANT III, 0 on IV) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pustula
 Sambucus 475

16 R IV+V less than 0.95 x HT II . . . . . . . . . . . . . . . . . . . . . . . . . . indica*

- R IV+V more than 0.95 x HT II . . . . . . . . . . . . . . . . . . . . . . hendersoni
17 Distal part of SIPH swollen to more than 1.7 x minimum diameter of
proximal part (Fig. 108I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . salicis
- Distal part of SIPH much less swollen, rarely more than 1.5 x minimum
diameter of proximal part (e.g. Fig. 108J) . . . . . . . . . . . . . . . . . . . . . . 18
18 Process on ABD TERG 8 very large and cowl-like, extending over tip of
CAUDA, tapering almost to a point from a very broad base, and bearing
3-5 additional short hairs besides the 2 hairs near the apex (Fig. 107I).
SIPH 2.4-2.7 x CAUDA, which is short and flask-shaped . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aspidaphoides
- Process on ABD TERG 8 usually smaller, if extending beyond tip of
CAUDA then the distal part of the process is finger-like, with a rounded
apex, and with only the 2 hairs near the apex. SIPH 1.6-2.6 x CAUDA,
which is tongue-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 SIPH dusky, 1.6-1.9 x CAUDA. Process on ABD TERG 8 short, about
0.5 x CAUDA or less, never broad-based (Fig. 107F) . . . . . . borealis
- SIPH (e.g. Fig. 108J) 1.9-2.6 x CAUDA. Process on ABD TERG 8 more
than 0.5 x CAUDA, and often extending finger-like from a broad base
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 CAUDA with 4 hairs, rarely with 5 in which case all are lateral. Process
on ABD TERG 8 often extending beyond tip of CAUDA (Fig. 107G).
(Alata with 5-9 secondary rhinaria on ANT IV and 3-4 on V) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rutila
- CAUDA almost always with 5 hairs, 4 on the sides and 1 dorsad. Process
on ABD TERG 8 rarely extending beyond tip of CAUDA (Fig. 107H).
(Alata with 0-2 secondary rhinaria on ANT IV and none on V . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aegopodii

SAMBUCUS Elders Caprifoliaceae

Most Sambucus species are shrubs; nevertheless a full list of elder aphids and
their recorded hosts is given for the sake of completeness.

Host Plant List

Sambucus australis Aphis sambuci
S, buergeriana Aphis horii
476 Sambucus

S. callicarpa Macrosiphum stanleyi

(Pacific Red Elder)
S, canadensis [Acyrthosiphon macrosiphum]
(American Elder) Aphis sambuci, spiraecola
Myzus persicae
S. cerulea Macrosiphum stanleyi
(Blue Elder) Myzus persicae
S. ebulus Aphis sambuci
S. melanocarpa Macrosiphum stanleyi
S. mexicana Myzus persicae
S. microbotrys Macrosiphum stanleyi
S. nigra Aphis sambuci
(European Black Elder)
S. pubens Macrosiphum stanleyi
(Scarlet Elder)
S. racemosa Aphis sambuci
(European Red Elder) Aulacorthum magnoliae
S. sieboldiana Aphis horii, sambuci
Aulacorthum magnoliae
S. thunbergiana Aphis sambuci
S. williamsii var. coreana Aphis horii
Aulacorthum magnoliae

Key to Elder aphids (apterae viviparae only)

1 Large pale aphid (BL more than 3 mm). SIPH long and pale, more than
3 x CAUDA, with distal swelling and a subapical zone of polygonal
reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum stanleyi
BL less than 3 mm. SIPH if pale less than 3 x CAUDA and/or without
subapical reticulation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Abdomen with marginal tubercles only on ABD TERG 1 and 7, or with
any marginal tubercles on other tergites small and inconsistently present.
SIPH pale or dark, but if dark and tapering then less than 2 x CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532
ABD TERG 1-4 (or 1-5) and 7 with large domed marginal tubercles
placed between the dark marginal (prespiracular) sclerites. SIPH dark,
tapering, 2.2-4.0 X CAUDA which is short, dark and blunt . . . . . . . 3
3 Hairs on ANT III long and very fine, the longest 1.7-2.1 x basal diameter
of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis horii
 Sasa 477

Hairs on ANT III shorter, the longest 0.5-1.7 x basal diameter of

segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis sambuci

SAPINDUS Sapindaceae

Sapindus mukorossi Tinocallis insularis

SAPIUM Euphorbiaceae

Sapium sebiferum Toxoptera odinae

(Chinese Tallow Tree)

SARCOCEPHALUS Rubiaceae

Sarcocephalus bartlingeri Sinomegoura citricola

SASA Bamboos Gramineae

Host Plant List

Sasa albomarginata Paracolopha morrisoni
S. nipponica Takecallis arundicolens, sasae
S. paniculata = S. senanensis
S. ramosa Ceratovacuna japonica
Paracolopha morrisoni
Takecallis arundicolens
S. senanensis Melanaphis bambusae
Paracolopha morrisoni
Takecallis arundicolens, sasae
Yezaphis sasicola
S. veitchii Paracolopha morrisoni
478 Sassafras

Compared with Bambusa and Arundinaria, the aphid fauna of Sasa seems
more limited. Use key to aphids on Arundinaria. Ceratovacuna japonica will
come out to C. sylvestris in that key, apart from the shorter frontal horns;
separation of Ceratovacuna spp. on bamboos is provided in the key to aphids
on Bambusa, couplets 27-29. Yezaphis sasicola Matsumura cannot be keyed
as no specimens are available (but see information from original description
summarized on p. 925).

SASSAFRAS Lauraceae

Sassafras albidum Euthoracaphis umbellulariae

SCHEFFLERA Araliaceae

Schefflera actinophylla Aphis hederae

S. octophylla Aulacorthum circumflexum
S. venulosa Myzus persicae

Key to aphids on Schefflera

- SIPH tapering and black like CAUDA which bears 9-18 hairs. R IV+V
1.35-1.60 X HT II. ANT PT/BASE 1.8-2.8 . . . . . . . . . . Aphis hederae
SIPH and CAUDA pale or other characters different . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

SCHIMA Theaceae

Schima noronhae Greenidea schimae

(Guger Tree)
S. wallichii Greenidea schimae
Schima sp. Greenidea schimae
 Sclerocarya 479

SCHINUS Peppertrees Anacardiaceae

Schinus dependens Aphis schinifoliae

S. motte Aphis gossypii, schinifoliae
S. terebinthifolius Aphis gossypii
Toxoptera aurantii

Key to aphids on Schinus

- SIPH pale. ANT PT/BASE less than 1.7 . . . . . . . . . Aphis schinifoliae
- SIPH dark. ANT PT/BASE more than 1.9 . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

SCHOTIA Leguminosae

Schotia brachypetala Myzus persicae

S. latifolia Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

SCHOUTENIA Tiliaceae

Schoutenia ovata Cervaphis schouteniae

SCLEROCARYA Anacardiaceae

Sclerocarya birrea Aphis gossypii

480 Scolopia

SCOLOPIA Flacourtiaceae

Scolopia crenata Toxoptera aurantii

S. ecklonii Myzus persicae
(Use key to polyphagous aphids, p. 532)

SECURINEGA Euphorbiaceae

Securinega virosa Schoutedenia ralumensis

(= Phyllanthus virosus) Sitobion sp.
Toxoptera citricidus
(These species are separated in the key to aphids on Bridelia, p. 102)

SEQUOIA Taxodiaceae

Sequoia sempervirens Cinara cupressi

Illinoia morrisoni

- ANT PT/BASE less than 0.5. SIPH as large dark hairy cones . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara cupressi
- ANT PT/BASE 3.7-4.4. SIPH tubular and swollen on distal half .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni

SEQUOIADENDRON Taxodiaceae

S. giganteum Illinoia morrisoni

 Sindora 481

SESBANIA Leguminosae

Host Plant List

Sesbania aculeata Aphis craccivora
S. aegyptiaca Aphis craccivora
S. coerulescens Aphis craccivora
S. grandiflora Acyrthosiphon gossypii
S. macrantha Sitobion sp.
S. punicea Aphis craccivora
S. sesban Aphis craccivora
Macrosiphum euphorbiae
Myzus persicae

Key to aphids on Sesbania

1 SIPH pale, long and thin, about 3 x CAUDA . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon gossypii
- SIPH about 2 x CAUDA or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 SIPH with distal band of polygonal reticulation and hairs on ANT III
less than 0.5 x basal diameter of segment . . . . . . . . . . . . . . Sitobion sp.
SIPH either without a distal band of polygonal reticulation or hairs on
ANT III more than 0.5 x basal diameter of segment . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

SHIIA = CASTANOPSIS

SINDORA Leguminosae

Sindora sp. Anomalosiphum murphyi

482 Sinojackia

SINOJACKIA Styracaceae

Sinojackia xylocarpa Aleurodaphis sp.

SMODINGIUM Anacardiaceae

Smodingium argutum Toxoptera odinae

SOPHORA Leguminosae

Sophora alopecuroides Acyrthosiphon sophorae

Aphis craccivora
S. japonica Acyrthosiphon gossypii, sophorae
(Pagoda Tree) Aphis sophoricola
Appendiseta robiniae
Tinocallis sophorae
S. moorcroftiana Myzus persicae

Key to aphids on Sophora

(Tentative, as host-specific species not seen by authors)
1 All viviparae alate. CAUDA knobbed, anal plate bilobed. Secondary
rhinaria on ANT III transversely elongate. ANT PT/BASE about 0.5.
SIPH short, truncated cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Viviparae apterous or alate. CAUDA tongue- or finger-shaped, anal plate
entire. ANT PT/BASE more than 2. SIPH tubular . . . . . . . . . . . . . . . 3
2 SIPH with a single hair appended at base. ABD TERG 1 and 2 without
finger-like tubercles . . . . . . . . . . . . . . . . . . . . . . . . . Appendiseta robiniae
- SIPH without an appended hair, ABD TERG 1 and 2 with paired finger-
like tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis sophorae*
3 CAUDA and SIPH pale, antennal tubercles well developed . . . . . . . . 4
CAUDA and SIPH black, antennal tubercles weakly developed . . . . 6
 Sorbus 483

4 Antennal tubercles with inner faces convergent, scabrous (Fig. 42I). SIPH
slightly swollen on distal half . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
Antennal tubercles divergent, smooth. SIPH tapering . . . . . . . . . . . . . 5
5 SIPH 0.33-0.50 x BL . . . . . . . . . . . . . . . . . . . . . Acyrthosiphon gossypii
SIPH about 0.3 x BL or less . . . . . . . . . . . . . Acyrthosiphon sophorae*
6 Abdominal dorsum with an extensive black patch . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
Abdominal dorsum with separate irregular spinal and small marginal
sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis sophoricola*

SORBUS Rowans, Whitebeams Rosaceae

Host Plant List

Sorbus alnifolia Nippolachnus piri
[Rhopalosiphum rufiabdominalis]
Sappaphis piri
S. americana Aphis pomi
(American Mountain Ash) Eriosoma lanigerum
Fimbriaphis gentneri
Muscaphis drepanosiphoides
Rhopalosiphum insertum
S. amurensis [Schizaphis sp. (Paik, 1972)]
S. aria Aphis lantanae
(Whitebeam) Dysaphis ariae
Myzus persicae
S. aucuparia Aphis spiraecola
(European Mountain Ash or Brachycaudus helichrysi
Rowan) Dysaphis [aucupariae], sorbi
Eriosoma sorbiradicis
Macrosiphum pyrifoliae
Muscaphis drepanosiphoides (incl.
ssp. irae)
Myzus ornatus
Nearctaphis californica
Ovatus insitus
Rhopalosiphum insertum
S. caucasigena Dysaphis sorbi
484 Sorbus

S. commixta [Myzaphis komatsubarae Shinji,

(Japanese Rowan) 1922c; possibly a Dysaphis?]
Prociphilus oriens
S. cuspidata Dysaphis pavlovskyana ssp. indica
S. domestica Dysaphis [aucupariae], sorbi
(Service Tree) Rhopalosiphum insertum
S. foliolosa Indotuberoaphis sorbi
S. graeca Dysaphis sp. near pavlovskyana
S. x hybrida Aphis pomi
Dysaphis sorbi
S. intermedia Aphis pomi
Dysaphis sorbi
Rhopalosiphum insertum
S. persica Dysaphis pavlovskyana
S. quercifolia Rhopalosiphum insertum
S. scopulina Nearctaphis yohoensis
S. sitchensis Macrosiphum pyrifoliae
Muscaphis drepanosiphoides
S. torminalis Dysaphis aucupariae
(Wild Service Tree)
S. turkestanica Dysaphis pavlovskyana
Sorbus spp. Prociphilus corrugatans
Sorbaphis chaetosiphon

Key to species on Sorbus (apterae viviparae except where otherwise stated)

1 ANT PT/BASE less than 0.5. CAUDA broadly rounded . . . . . . . . . . 2
- ANT PT/BASE more than 0.5. CAUDA semicircular, helmet-, tongue-
or finger-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2 Wax glands absent. Eyes multifaceted, without an ocular tubercle. Body
pale, elongate, more than 2 x longer than wide. (Alata with a quadrate
black dorsal abdominal patch) . . . . . . . . . . . . . . . . . . . Nippolachnus piri
- Wax glands present. Eyes 3-faceted. Body broad, less than 2 x longer
than wide. (Alata without dorsal abdominal pigment) . . . . . . . . . . . . . 3
3 SIPH present as pores with surrounding hairs. Wax pore plates each
comprising a ring or group of facets around one or more clear central
areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
- SIPH absent or inconspicuous. Wax pore plates honeycomb-like, without
clear central areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
 Sorbus 485

Fig. 109. A, wax pore plate of Eriosoma sorbiradicis (aptera); section of dorsal cuticle (ABD TERG 3) of
B, Muscaphis drepanosiphoides (aptera), C, Nearctaphis yohoensis (aptera); D, SIPH of Dysaphis pavlovskyana.

4 BL more than 1.5mm. Total antennal length 0.29-0.46 mm. Wax pore
plates with central areas of irregular shape, often subdivided (Fig. 60C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanigerum
- BL less than 1.4 mm, with total antennal length only 0.12-0.18 mm. Wax
pore plates with always undivided, approximately circular, central areas
(Fig. 109A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma sorbiradicis
5 R IV+V (of fundatrix) more than 0.2mm long and bearing 7 or more
accessory hairs. BL of spring migrant alata more than 4mm . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
- R IV+V (of fundatrix) less than 0.2 mm long with 2-4 accessory hairs. BL
of spring migrant alata less than 3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus corrugatans
6 CAUDA short; helmet-shaped, triangular or semicircular, shorter than or
about as long as its basal width in dorsal view . . . . . . . . . . . . . . . . . . . 7
- CAUDA tongue- or finger-shaped, clearly longer than its basal width
in dorsal view . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
7 Dorsal cuticle of aptera (fundatrix) very rough with numerous small
denticulate projections, and with fine, long hairs (Fig. 109B). Alata with
transverse bars of dorsal abdominal pigment, often broken, never fused
between tergites . . . . . . . . . . . . . . . . . . . . . . Muscaphis drepanosiphoides
- Dorsal cuticle of aptera (fundatrix or subsequent generation) smooth
486 Sorbus

or weakly reticulate, with long or short hairs. Alata with a solid black
dorsal abdominal patch, or with extensive bars fused between some
tergites ...................................................... 8
8 SIPH more than 4 x CAUDA and bearing a number of long hairs (in both
fundatrix and spring migrant alata) . . . . . . . . . Sorbaphis chaetosiphon
- SIPH less than 4 x CAUDA and without hairs . . . . . . . . . . . . . . . . . . 9
9 SIPH with closely-spaced rows of imbrications bearing small nodules or
spinules (e.g. Fig. 42M) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- SIPH smooth or normally imbricated, the imbrications sometimes spinu-
lose but not in regular closely-spaced rows . . . . . . . . . . . . . . . . . . . . . 11
10 ABD TERG 2-4 each with incomplete double rows of 6-10 spinopleural
hairs 50-70 µm long with fine apices. Tergum rough, with numerous
spinules arranged on a distinct reticulation (Fig. 109C), and with an
extensive solid black abdominal patch . . . . . . . . Nearctaphis yohoensis
- ABD TERG 2-4 each with a single row of 4 spinopleural hairs that are
blunt or acuminate. Tergum with faint reticulation and few spinules, and
broad transverse sclerotic bars that are not or only partially fused between
tergites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nearctaphis californica
11 SIPH smooth, less than 2 x width at base. Spiracular apertures rounded
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachycaudus helichrysi
- SIPH imbricated, more than 2 x width at base. Spiracular apertures small
and reniform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 SIPH pale, about 2 x CAUDA or less, and distinctly swollen on distal
half (Fig. 109D) . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis pavlovskyana
- SIPH pale or dark, more than 2 x CAUDA, cylindrical or tapering or
only very slightly swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 SIPH pale. Marginal tubercles present on ABD TERG 1-7 (sometimes
absent from 6). CAUDA with 6-12 hairs . . . . . . . . . . . . Dysaphis sorbi
- SIPH dark, at least on distal part. Marginal tubercles present on ABD
TERG 1-4 or 1-5, absent from 6 and 7. CAUDA with 5, or rarely 6,
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14 Hairs on ANT III pointed, maximally 0.9-1.5 x basal diameter of
segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis aucupariae
- Hairs on ANT III blunt, maximally 0.3-0.8 x basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Hairs on ABD TERG 1-3 consistently short and blunt, maximally
15-20 µm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis ariae
- Hairs on ABD TERG 1-3 of variable length, often short and blunt
but usually with at least some pointed hairs 20-60 µm long, especially
marginally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dysaphis plantaginea
 Sorbus 487

16 ANT PT/BASE 0.55-0.66. Body with finger-like dorsal processes (head

with 2 pairs, thoracic segments and ABD TERG 1-4 each with one spinal
and one marginal pair, and ABD TERG 5-8 each with a single medial
process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Indotuberoaphis sorbi
- ANT PT/BASE more than 1. Body without finger-like dorsal processes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 Antennal tubercles undeveloped or weakly developed, not exceeding
height of central part of frons in dorsal view. CAUDA and anal plate
dusky to dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
- Antennal tubercles moderately to well developed, clearly exceeding height
of central part of frons in dorsal view. CAUDA and anal plate pale or
only slightly dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
18 SIPH slightly swollen or cylindrical over most of length, constricted
subapically and with a large flange (Fig. 60H). Marginal tubercles on
ABD TERG 7 placed posteriodorsad of spiracle and with basal diameter
usually smaller than spiracular aperture . . . . Rhopalosiphum insertum
- SIPH tapering from base to flange, which is only moderately developed.
Marginal tubercles on ABD TERG 7 placed posterioventrad of spiracle,
and usually larger than spiracular aperture . . . . . . . . . . . . . . . . . . . . . 19
19 Longest hairs on ANT III more than 2 x basal diameter of segment.
ANT PT/BASE more than 3. Alata with secondary rhinaria distributed
ANT III 19-28, IV 1-11, V 0-3. CAUDA tapering, less than 2 x its
basal width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis lantanae
- Longest hairs on ANT III less than 1.8 x basal diameter of segment.
ANT PT/BASE less than 3. Alata with secondary rhinaria distributed
ANT III 6-11, IV 0-4, V 0. CAUDA finger-like, more than 2 x its basal
width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20 Marginal tubercles present on ABD TERG 2-4. CAUDA with 10-19
hairs (rarely less than 13). R IV+V more than 0.13 mm . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis pomi
- Marginal tubercles absent from ABD TERG 2-4. CAUDA with 7-15 hairs
(rarely more than 12). R IV+V less than 0.125 . . . . . Aphis spiraecola
21 Inner faces of antennal tubercles scabrous and convergent in dorsal view
(e.g. Fig.42I) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
- Inner faces of antennal tubercles smooth or nearly so, and divergent in
dorsal view (e.g. Fig. 42K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
22 Dorsal abdomen with a pattern of dark intersegmental spots and flecks
(Fig. 121J). ANT PT/BASE less than 2.5 . . . . . . . . . . . . Myzus ornatus
- Dorsal abdomen without clear intersegmental dark markings. ANT PT/
BASE more than 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
488 Spathodea

23 SIPH slightly swollen on distal half (Fig. 121H). ANT PT/BASE less
than 5. (Alata with secondary rhinaria only on ANT III) . . . . . . . . . . .
................................................. Myzus persicae
- SIPH tapering or cylindrical on distal half. ANT PT/BASE more than
5. (Alata with secondary rhinaria distributed ANT III 60-83, IV 36-52,
V 13-22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovatus insitus
24 Body spindle-shaped. SIPH with a subapical zone of polygonal reticula-
tion (Fig. 121C). ANT III usually with several secondary rhinaria on basal
half, and with longest hairs more than half basal diameter of segment
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae/pyrifoliae
- Body oval. SIPH without any polygonal reticulation (Fig. 42Q). ANT
III with one small rhinarium near base, and with minute hairs of length
less than half basal diameter of segment . . . . . . . Fimbriaphis gentneri

SPATHODEA Bignoniaceae

Spathodea campanulata Aphis gossypii, spiraecola

S. diepenhorstii Toxoptera odinae
(Use key to polyphagous aphids, p. 532)

SPONDIAS Anacardiaceae

Spondias cirouella Aphis spiraecola

S. mombin Aphis spiraecola
Toxoptera aurantii
S. purpurea Aphis spiraecola
Myzus persicae
(Use key to polyphagous aphids, p. 532)

STAPHYLEA Staphyleaceae

Staphylea bumalda Indomegoura indica, nigrotibiae

S. pinnata Rhopalosiphoninus staphyleae
 Staphylea 489

Fig. 110. SIPH of A, Indomegoura indica, B, /. nigrotibiae and C, Rhopalosiphoninus staphyleae.

Key to aphids on Staphylea

1 Legs mainly pale. Head spiculose, with inner faces of antennal tubercles
approximately parallel. SIPH pale except distally, swollen on distal 0.7,
without subapical reticulation; maximum diameter of swollen part more
than 1.5 x minimum width of proximal 'neck' region (Fig. 110C) ....
.................................... Rhopalosiphoninus staphyleae
Legs dark. Cuticle of head smooth or slightly wrinkled, inner faces of
antennal tubercles broadly divergent. SIPH uniformly dark, more-or-less
barrel-shaped, thickest in middle, with subapical reticulation (Figs 110A,
B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 BL more than 3 mm. CAUDA elongate, much longer than wide, more
than 0.5 x SIPH. R IV+V 0.7-0.9 x HT II. SIPH almost cylindrical on
proximal 0.7 of length, narrowing only subapically (Fig. 110A) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Indomegoura indica
BL less than 3 mm. CAUDA triangular, about as long as its basal width,
less than 0.5 x SIPH. R IV+V 0.9-1.1 x HT II. SIPH narrowing prox-
imally as well as distally (Fig. 110B) . . . . . . . . Indomegoura nigrotibiae
490 Steganotaenia

STEGANOTAENIA Umbelliferae

Steganotaenia araliacea Hydaphis coriandri (see B & E,

1984:280-281)

STEWARTIA = STUARTIA

STREBLUS Moraceae

Streblus sp. Greenideoida hannae

STRYCHNOS Loganiaceae

Strychnos spinosa Aphis gossypii

S. pseudoguina Aphis gossypii

STUARTIA Theaceae

Stuartia sinensis Myzus persicae

STYRAX Styracaceae

The life cycles of most Hormaphidinae described from galls on Styrax are
not yet known, although some have recently been elucidated. Most species
were originally described in Astegopteryx, but as the secondary host forms
became known some have been transferred to other genera (e.g. Ceratovacuna
 Styrax 491

nekoashi and Ceratoglyphina styracicola), or synonymized with known species

on secondary hosts (e.g. A. shitosanensis = Pseudoregma koshunensis). At
present generic concepts within the group are hazy, but it seems likely that
Astegopteryx and Ceratoglyphina alternate between Styrax and bamboos,
whereas other genera may have developed secondary host associations with a
wider range of Gramineae.
The key which follows the host plant list is somewhat tentative. Any
identification should be checked against the description of the gall of the
species given in Section III of the book (see also Fig. 124).

Host Plant List

Styrax benzoin Astegopteryx lambersi, setigera,
styracophila
Cerataphis fransseni
Ceratoglyphina bambusae
S. ferrugineum Toxoptera aurantii
S. formosanum Ceratoglyphina styracicola
Ceratovacuna nekoashi
Tuberaphis taiwana
S. japonica Aphis craccivora
(Snowball Tree) Ceratovacuna japonica, nekoashi
Toxoptera aurantii
Tuberaphis takenouchii
S. hookeri 'gall of unknown aphid' - Mani,
1973: 194
S. obassia Aphis fabae
Ceratoglyphina bambusae
Ceratovacuna nekoashi
Hamiltonaphis styraci
S. officinalis Toxoptera aurantii
S. paralleloneurum Astegopteryx roepkei
S. serrulatum Astegopteryx leeuweni, sumatrana
(var. mollissimus) Tuberaphis vandermeermohri
S. subdenticulatum Astegopteryx sumatrana
Tuberaphis vandermeermohri
S. suberifolia Astegopteryx bambucifoliae,
jamaritsu,
Ceratoglyphina styracicola
Ceratovacuna nekoashi
Pseudoregma bambucicola,
koshunensis
Toxoptera aurantii
492 Styrax

Styrax spp. Astegopteryx chinensis

Astegopteryx sp. (Thailand)

Tentative key to alatae of Hormaphidinae from galls on Styrax

(based partly on Takahashi, 1936b)
Check characters for group: alatae have 5-segmented antennae with short PT
and annular secondary rhinaria, combined with a once-branched media in the
forewing. Otherwise try key to polyphagous aphids, p. 532.
1 ANT IV and V without secondary rhinaria (rarely IV with 1) . . . . . . 2
- ANT IV usually with more than 1 secondary rhinarium, V with secondary
rhinaria present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 R IV+V longer than ANT IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- R IV+V shorter than ANT IV. (ANT III with 10-16 secondary rhinaria)
............................................................. 4
3 ANT III with 0-14 secondary rhinaria. ANT IV more than 0.75 X ANT
V (incl. PT) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx jamaritsu
- ANT III with 17-25 secondary rhinaria. ANT IV less than 0.75 x ANT
V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx sumatrana
4 ANT IV less than 0.75 x ANT V (incl. PT). The 2 outer hairs on first
segments of fore and mid-tarsi very long and fine, about 2.5 x longer than
medial hairs (Fig. 111A) . . . . . . Tuberaphis taiwana/vandermeermohri
- ANT IV more than 0.9 x ANT V (incl. PT). The 2 outer hairs on first
segments of fore and mid-tarsi about 1.5 x longer than medial hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx lambersi
5 ANT III distinctly longer than (1.2-1.6 x) IV+V (incl. PT) together
............................................................. 6
- ANT III shorter than, about equal to, or up to 1.15 x longer than, IV+V
together . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6 ANT V with more than 11 secondary rhinaria . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx chinensis*
- ANT V with 0-11 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 ANT III much thicker than IV and bearing 28-39 closely-spaced secon-
dary rhinaria (Fig. 111B) . . . . . . . . . . . . . . . . . . . . Astegopteryx leeuweni
- ANT III similar in thickness to IV and bearing 15-25 secondary rhinaria
............................................................. 8
8 Front of head with 32-40 small hairs, extending across area immediately
posteriodorsal to median ocellus. ANT V with 6-11 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Astegopteryx setigera*
 Styrax 493

Fig. 111. A, fore tarsus of Tuberaphis vandermeermohri (alata from gall); B, antenna of alata from gall
of Astegopteryx leeuweni; C, ANT V of alata from gall of Cerataphis fransseni; D, ANT V of alata from
gall of Ceratoglyphina styracicola.

- Front of head with less than 20 hairs, none of which are immediately
anteriodorsal to median ocellus. ANT V with 0-8 secondary rhinaria
............................................................. 9
9 ANT V with 0-4 secondary rhinaria, and with primary rhinarium extend-
ing across more than half width of segment (Fig. 111C) . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cerataphis fransseni
- ANT V with 4-8 secondary rhinaria, and with a very small primary
rhinarium extending across less than half width of segment (Fig. 111D)
....................................... Ceratoglyphina styracicola
10 ANT V usually without secondary rhinaria, rarely with 1-2 . . . . . . . . .
.......................................... Tuberaphis takenouchii
- ANT V with 3-16 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11 ANT V with 10-16 secondary rhinaria. SIPH without any clearly asso-
ciated hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratovacuna nekoashi
- ANT V with 3-11 secondary rhinaria. SIPH with 2-7 hairs around it
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 ABD TERG 7-8 distinctly sclerotized, thickened and dusky. (Secondary
494 Swietenia

rhinaria distributed ANT III 20-27, IV 8-15, V 6-11) . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoregma koshunensis/bambucicola
- ABD TERG 7-8 not thickened, only sclerotized around hair bases if at
all . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13 Hairs on ABD TERG 7-8 with small basal sclerites. (Secondary rhinaria
distributed III 18-26, IV 7-10, V 6-9) . . . . . . . . . Astegopteryx roepkei
- Hairs on ABD TERG 7-8 without basal sclerites . . . . . . . . . . . . . . . . 14
14 Head with small irrregular pale dorsomedial areas. (Secondary rhinaria
distributed III 18-20, IV 6-9, V 4-8) .... Astegopteryx bambucifoliae
- Head without paler dorsomedial areas . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 ANT III clearly thicker and less than 1.3 x longer than ANT IV. (Secon-
dary rhinaria distributed III 10-15, IV 7-11, V 4-7) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceratovacuna japonica*
- ANT III similar in thickness to and more than 1.6 x longer than ANT
IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16 ANT PT distinctly longer than its basal width. Embryos without frontal
horns (sexuparae). Secondary rhinaria distributed III 12-19, IV 5-10,
V 3-9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hamiltonaphis styraci *
- ANT PT only about as long as its basal width or shorter. Embryos with
frontal horns visible through maternal cuticle. Secondary rhinaria dis-
tributed III 17-25, IV 6-14, V 6-9 . . . . . . . . Astegopteryx styracophila

SWIETENIA Mahoganies Meliaceae

No aphids have been recorded.

SYMPLOCOS Symplocaceae

Symplocos laurina Greenidea symplocosis

S. sessilifolia Sinomegoura symplocois
Toxoptera aurantii
Symplocos sp. [Micromyzus sp. (Basu, Ghosh
and Raychaudhuri, 1970)]
 Syringa 495

Key to aphids on Symplocos

1 SIPH with numerous long hairs . . . . . . . . . . . . . Greenidea symplocosis
- SIPH without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 CAUDA long, pointed, darker than SIPH, with about 8 hairs. Antennal
tubercles well developed. Hairs on ANT III less than half basal diameter
of segment. No stridulatory apparatus . . . . . . Sinomegoura symplocois
SIPH as dark as CAUDA which is blunt and bears more than 10 hairs.
Antennal tubercles weakly developed. Longest hairs on ANT III more
than half basal diameter of segment. Stridulatory ridges present on
abdominal sternites 5-6 (Fig. 122) . . . . . . . . . . . . . . . Toxoptera aurantii
(or use key to polyphagous aphids on p. 532)

SYNCARPIA Turpentines Myrtaceae

Syncarpia laurifolia Aphis gossypii

SYRINGA Oleaceae

Host Plant List

Syringa amurensis Aulacorthum syringae
Prociphilus oriens
S. emodi Aulacorthum syringae
Prociphilus oriens
S. japonica Aulacorthum syringae
Prociphilus oriens
S. persica Aphis craccivora
S, reticulata Aulacorthum syringae
S. vulgaris Aulacorthum syringae
(Lilac) Myzus persicae
Prociphilus americanus, bumeliae

Key to aphids on Syringa

1 Wax glands present (especially obvious in aptera as large clear areas on
head). ANT PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
496 Syzygium

- Wax glands absent. ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . 3

2 Alata with at least 4 secondary rhinaria on ANT V . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
Alata with 0-2 secondary rhinaria on ANT V . . . . . . . . . . . . . . . . . . . . 3
3 ANT III more than 5 x ANT II, as long as or longer than ANT IV+V
together. Head with a pair of very large posterior dorsal wax pore plates,
conspicuous as clearly delimited pale areas . . . . . Prociphilus bumeliae
- ANT III less than 5 x ANT II, usually shorter than ANT IV+V together.
Head without or with only small wax pore plates, not forming clearly
delimited pale areas . . . . . . . . . . . . . . . . . . . . . . . Prociphilus americanus
4 ANT III-VI of aptera dark, with PT/BASE more than 5. Tibiae wholly
dark, femora contrastingly pale except at apices. SIPH dark, tapering,
with a well-developed flange. CAUDA pale . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum syringae
Without this combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

SYZYGIUM Myrtaceae

Host Plant List

Syzygium antisepticum [Eutrichosiphum sp. (Noordam, in
prep.)]
[Greenidea sp. (Noordam, in
prep.)]
[Mollitrichosiphum sp. (Noordam,
in prep.)]
S. buxifolium Taiwanaphis decaspermi
S. lineatum [Greenidea sp. (Noordam, in
prep.)]
Taiwanaphis atuberculata
S. racemosum [Greenidea sp. (Noordam, in
prep.)]
Taiwanaphis montanicola
S. syzygioides [Greenidea sp. (Noordam, in
prep.)]
Taiwanaphis pseudocaudata
Syzygium sp. (?) Taiwanaphis atrovirens
 Tabebuia 497

Key to aphids on Syzygium

1 SIPH long, tubular, with numerous long hairs . . . . Greenideinae spp.
[Descriptions of a Greenidea sp., a Eutrichosiphum sp. and a Mollitri-
chosiphum sp. from Java are in preparation (Noordam, pers. comm.)]
SIPH short, broad-based, conical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT III 2.1-2.9 x PT. Fourth instar nymph with an unpaired scabrous
median process on ABD TERG 8, but without paired lateral processes
(Fig. 112A) . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwanaphis pseudocaudata
ANT III 3.1-7.0 x PT. Fourth instar nymph without a single median pro-
cess on ABD TERG 8, with or without paired lateral processes . . . . 3
3 BL 2.0-2.1 x total length of antenna, which is 1.9-2.1 x head width
across (and including) eyes. Fourth instar nymph (and aptera) without
paired lateral processes . . . . . . . . . . . . . . . . . Taiwanaphis atuberculata*
- BL 1.2-1.6 x total length of antenna, which is 2.2-3.2 x head width
across eyes. Fourth instar nymph with or without paired lateral processes
............................................................. 4
4 ANT PT/BASE 0.8-1.15. ANT III 3.1-3.8 X PT. Fourth instar nymph
with paired lateral processes on ABD TERG 6-8 (plus a subdivided medial
process on ABD TERG 8; Fig. 112B) . . . . . . . Taiwanaphis atrovirens*
- ANT PT/BASE 0.4-0.7. ANT III 4.1-7.0 x PT. Fourth instar nymph
with or without paired lateral processes . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 ANT III of alata pale, and bearing 11-18 secondary rhinaria. Fourth
instar nymph without paired lateral processes Taiwanaphis decaspermi
ANT III of alata dark, bearing 19-41 secondary rhinaria. Fourth instar
nymph with paired lateral processes (Fig. 112C) . . . . . . . . Taiwanaphis
montanicola

TABEBUIA Bignoniaceae

Tabebuia avellanedae Myzus persicae

T. chrysotricha Myzus persicae
T. heterophylla = T. pallida
T. pallida Aphis craccivora, gossypii
T. rosea Myzus persicae
(Use key to polyphagous aphids, p. 532)
498 Tabernaemontana

Fig. 112. End of abdomen of fourth instar nymph of A, Taiwanaphis pseudocaudata, B, T. atrovirens
and C, T. montanicola (all after Noordam and Hille Ris Lambers, 1985).

TABERNAEMONTANA Apocynaceae

Tabernaemontana chippii Sitobion krahi

T. coronaria Myzus persicae

TAMARINDUS Leguminosae

Tamarindus indica Aphis craccivora

TAMARIX Tamaricaceae

Host Plant List

Tamarix africana Brachyunguis tamaricis
 Tamarix 499

T. aphylla Brachyunguis tamaricophila

Myzus persicae
Toxoptera aurantii
T. articulata Brachyunguis tamaricophila
T. gallica Brachyunguis tamaricis
Brevicorynella quadrimaculata
T. karakalensis Brachyunguis tamaricophila
T. kotschyi Brachyunguis tamaricis
T. meyersi Brachyunguis tamaricophila
T. pallasii Brachyunguis tamaricis
T. ramosissima [Brachyunguis tamaricivorum
Narzikulov, 1954]
Brevicorynella quadrimaculata
Protrama tamaricis
Tamarix spp. Aphis craccivora, fabae,
viridissima

Key to aphids on Tamarix (apterous viviparae)

1 ANT PT/BASE more than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- ANT PT/BASE less than 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 SIPH and CAUDA pale or dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
SIPH and CAUDA black . . . go to key to polyphagous aphids, p. 532
(Aphis craccivora, A. fabae, Toxoptera aurantii)
3 Antennal tubercles undeveloped. SIPH tapering, about 1.5 x CAUDA
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis viridissima
Antennal tubercles well developed, their inner faces convergent in dorsal
view (Fig. 42I). SIPH slightly swollen distally, about 2 x CAUDA (Fig.
121H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
4 BL more than 3.5mm. HT II much elongated. SIPH pore-like, on very
flat pigmented cones . . . . . . . . . . . . . . . . . . . . . . . . . . Protrama tamaricis
BL less than 2mm. HT II normal. SIPH not pore-like . . . . . . . . . . . 5
5 Eye without an ocular tubercle. SIPH slightly swollen in middle. Dorsum
with 2 pairs of black spots . . . . . . . . . . Brevicorynella quadrimaculata*
Eye with normal 3-faceted ocular tubercle. SIPH tapering. Dorsum
without dark markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Antenna 6-segmented, with ANT III much longer than ANT VI (incl. PT).
CAUDA triangular, 1.7-2.3 x SIPH, which are 1.0-1.3 x their basal
500 Taxodium

Fig. 113. End of abdomen of aptera of A, Brachyunguis tamaricophila and B, B. tamaricis.

widths (Fig. 113A). R IV+V 0.8-0.9 x HT II . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyunguis tamaricophila
Antenna 5- or 6-segmented, when 6-segmented ANT III is about as long
as ANT VI or shorter. CAUDA digitiform, 1.0-1.6 x SIPH, which
are 1.3-2.2 x their basal widths (Fig. 113B). R IV+V 0.6-0.75 x HT II
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brachyunguis tamaricis

TAXODIUM Taxodiaceae

Taxodium distichum Illinoia morrisoni

TAXUS Yews Taxaceae

Taxus baccata Prociphilus taxus

T. cuspidata Elatobium momii

- SIPH absent. R IV+V 7.0-8.0 x ANT PT . . . . . . . . . Prociphilus taxus

- SIPH tubular. R IV+V 0.2-0.4 x ANT PT . . . . . . . . Elatobium momii
 Terminalia 501

TECLEA Rutaceae

Tedea sp. Toxoptera citricidus

TECOMA Bignoniaceae

Tecoma capensis Aphis gossypii

Myzus persicae
T. radicans Aphis nerii
T. shirensis Myzus persicae
T. stans Aphis gossypii
Myzus ornatus, persicae
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

TECTONA Verbenaceae

Tectona grandis Aphis gossypii

Myzus persicae
(Use key to polyphagous aphids, p. 532)

TERMINALIA Combretaceae

Host Plant List

Terminalia arjuna Aphis gossypii
Paoliella nirmalae
T. australis Lizerius brasiliensis
T. brachystemma Aphis gossypii
Paoliella harteni, terminaliae
502 Terminalia

T. chebula 'gall of unknown aphid' - Mani,

1973: 143
T. mantaly Aphis gossypii
T. sericea Paoliella harteni, terminaliae

Key to aphids on Terminalia

1 CAUDA not knobbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
(or try key to polyphagous aphids, p. 532)
CAUDA knobbed (body of aptera, and sometimes alata also, with tubular
or wart-like dorsal processes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Knob of CAUDA very elongate, much longer than its maximal width,
tapering to apex (Fig. 114A) . . . . . . . . . . . . . . . . . . . Lizerius braziliensis
Knob of CAUDA about as long as its maximal width, with rounded apex
............................................................. 3
3 Adult viviparae apterous as well as alate. Alata with long and tapering
or finger-like spinal processes (Fig. 114B), like those of aptera, as well
as wart-like processes. Pronotum with 1 pair of anteriolateral hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paoliella nirmalae
Adult viviparae all alate. Spinal processes if present all short and wart-
like. Pronotum with 2 pairs of anteriolateral hairs . . . . . . . . . . . . . . . 4
4 Alata with paired, wart-like spinal hair-bearing processes on ABD TERG
1-8 (Fig. 114B), and similar marginal processes on ABD TERG 1-4. R
IV+V less than 0.75 x HT II . . . . . . . . . . . . . . . . . . . . . Paoliella harteni
Alata without abdominal processes. R IV+V more than 0.8 x HT II
............................................ Paoliella terminaliae

Fig. 114. A, CAUDA of Lizerius brasiliensis (aptera); B, examples of tapering and wart-like spinal
processes of Paoliella.
 Tilia 503

TETRACLINIS Cupressaceae

Tetraclinis articulata Cinara cupressi

THESPESIA Malvaceae

Thespesia acutiloba Aphis gossypii

THUJA Cupressaceae

Thuja occidentalis Cinara cupressi, louisianensis,

tujafilina
Th. orientalis Cinara louisianensis, tujafilina
Th. plicata Illinoia morrisoni
[Myzus persicae]
(Use key to aphids on Cupressus, p. 161)

TILIA Basswood, Lime, Linden Tiliaceae

Tilia americana Aulacorthum solani

(American Lime, Basswood) Eucallipterus tiliae
Longistigma caryae
Macrosiphum tiliae
Tiliphagus lycoposugus
T. amurensis Patchiella reaumuri ssp. orientalis
(Korean Lime) Tiliaphis coreana, shinae, shinjii
T. cordata Eucallipterus tiliae
(Small-leafed Lime) Patchiella reaumuri
T. x europea Eucallipterus tiliae
(Common Lime, Linden) Patchiella reaumuri
504 Tilia

T. insularis Tiliaphis shinae

T. japonica [Eucallipterus tilicola Shinji, 1933]
Rhopalosiphoninus tiliae
Tiliaphis shinae, shinjii
T. mandshurica Tiliaphis coreana, shinae
T. maximowicziana Prociphilus oriens
Rhopalosiphoninus tiliae
Tiliaphis shinae
T. miqueliana [Eucallipterus tilicola Shinji, 1933]
Rhopalosiphoninus tiliae
Tiliaphis shinae
T. orientalis Tiliaphis coreana
T. pekinensis Tiliaphis coreana, shinae
T. petiolaris Eucallipterus tiliae
T. platyphyllos. Eucallipterus tiliae
(Large-leafed Lime) Patchiella reaumuri
T. rubra Eucallipterus tiliae
T. tomentosa Eucallipterus tiliae
Patchiella reaumuri
T. tuan Shivaphis tilisucta
Tilia sp. Tiliaphis pseudoshinae

Key to aphids on Tilia (a revised and expanded version of the key in B & E,
1984)
1 SIPH absent or present only as inconspicuous pores (forming leaf-nest
galls) ........................................................ 2
- SIPH present either as short truncate cones, large pores on broad hairy
cones, or long tubes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2 Alata from gall with numerous elongate secondary rhinaria on ANT
III-V, not in a single row, protruding as sharp transverse ridges so that
the outline is serrate (Fig. 115A) . . . . . . . . . . . . Tiliphagus lycoposugus
- Alata from gall with secondary rhinaria on ANT III-V arranged in a row
and not protruding as sharp ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 BL of alata from gall more than 4 mm. Media of forewing unbranched.
Antenna with sparse hairs and at least 4 secondary rhinaria on ANT V
.............................................. Prociphilus oriens
- BL of alata from gall less than 3 mm. Media of forewing once-branched.
Antenna with numerous hairs and 0-3 secondary rhinaria on ANT V
(Fig. 115B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Patchiella reaumuri
 Tilia 505

Fig. 115. Antenna of spring migrant alata of A, Tiliphagus lycoposugus and B, Patchiella reaumuri.

4 BL more than 5 mm. Body and appendages covered in long hairs. SIPH
as large pores on broad hairy cones. (Alata with pterostigma extending
to tip of wing) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- BL less than 4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 SIPH tubular, much longer than their basal width. CAUDA finger-like
or triangular, anal plate entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- SIPH as truncate cones, hardly longer than their basal width. CAUDA
knobbed, anal plate bilobed (all viviparae alate) . . . . . . . . . . . . . . . . . . 8
6 SIPH black, markedly swollen over distal 0.7 of length . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhopalosiphoninus tiliae
- SIPH pale or dark, not at all swollen . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Aptera with SIPH mainly dark, about 3 x CAUDA, with a subapical
zone of polygonal reticulation; cuticle of head smooth; ANT III with 3-10
secondary rhinaria in a row . . . . . . . . . . . . . . . . . . . . Macrosiphum tiliae
- Aptera with SIPH pale except at apex, about 2 x CAUDA, without
polygonal reticulation; cuticle of head spiculose; ANT III with one small
secondary rhinarium near base . . . . . . . . . . . . . . . . . Aulacorthum solani
8 Secondary rhinaria on ANT III round to oval, with ciliated borders.
Forewing with Rs absent or indistinct and costal margin not heavily
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Shivaphis tilisucta
- Secondary rhinaria on ANT III transversely elongate, not ciliated. Costal
margin of forewing heavily pigmented and Rs present . . . . . . . . . . . . . 9
9 Forewing with separate fuscous patches at ends of veins. Hind femur
black except at base. ABD TERG 1-7 with black marginal as well as
pleural pairs of sclerites, and dark SIPH . . . . . . . . . Eucallipterus tiliae
- Forewing with a zig-zag black band joining distal ends of media, Cu la
and Cu lb . Hind femur mainly pale. ABD TERG 1-7 with or without
dark markings. SIPH pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
506 Tocoyena

10 Mesothoracic dorsal lobes each with a longitudinal dark stripe. Base of

forewing infuscated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- Mesothoracic dorsal lobes without longitudinal stripes. Base of forewing
clear ....................................................... 12
11 R IV+V and SIPH both shorter than HT II. ANT III with 19-33 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tiliaphis coreana
- R IV+V and SIPH both longer than HT II. ANT III with 8-16 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tiliaphis shinjii
12 R IV+V less than 0.8 x HT II. Embryos with very short spinal hairs
(about 4 µm long) on ABD TERG 1-6 . . . . . . . Tiliaphis pseudoshinae
- R IV+V 1.0-1.2 x HT II. Embryos with all spinal hairs long (35-40µm)
................................................ Tiliaphis shinae

TOCOYENA Rubiaceae

Tocoyena longiflora Aphis gossypii, spiraecola

(Use key to polyphagous aphids, p. 532)

TODDALIA Rutaceae

Toddalia asiatica Toxoptera aurantii

TOONA Meliaceae

Toona citiata Forda sp. (orientalis?)

TORRUBIA = PISONIA
 Trochodendron 507

TOURNEFORTIA Boraginaceae

Tournefortia hirsutissima Aphis gossypii

TREMA Ulmaceae

Trema amboinensis Toxoptera aurantii

T. orientalis Sumatraphis celti

SIPH tubular with distal ring of about 4 hairs and 1-2 hairs on the
basal half. ABD TERG 7 with 4 finger-like processes. ANT PT/BASE
0.25-0.30 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sumatraphis celti
Without this combination of characters . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

TRISTANIA Myrtaceae

Tristania conferta Aphis gossypii

Myzus ornatus
Toxoptera aurantii
(Use key to polyphagous aphids, p. 532)

TROCHODENDRON Trochodendraceae

Trochodendron aralioides Elatobium trochodendri

508 Tsuga

TSUGA Hemlocks Pinaceae

Host Plant List

Tsuga brunoniana Prociphilus taxus
T. canadensis Adelges funitecta
Prociphilus pini
T. chinensis Adelges funitecta
T. formosana Cinara taiwana
T. heterophylla Adelges funitecta
[Aphis fabae]
Cinara pilicornis, tsugae
Illinoia patriciae
T. mertensiana [Cinara wahsugae]
T. sieboldii Adelges funitecta
Mindarus ?abietinus
Tsugaphis sorini

Key to aphids on Tsuga (apterous parthenogenetic morph except where other-

wise stated)
1 Body pear-shaped, broadest anteriorly, with conspicuous wax pores on all
segments. Chitinous ovipositor present. Antennae very short, 3-segmented
............................................... Adelges funitecta
Body not broadest anteriorly, with or without wax glands. No chitinous
ovipositor. Antennae with 4-6 segments . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Body aleyrodiform; flattened, elliptical, divided about equally into
cephalothorax and abdomen, with legs and 4-segmented antennae con-
cealed beneath (e.g. Fig. 65B). CAUDA knobbed, anal plate bilobed .
................................................ Tsugaphis sorini
Body of normal aphid form, with exposed legs and 5- or 6-segmented
antennae. CAUDA variously shaped, anal plate entire . . . . . . . . . . . . 3
3 SIPH absent, or present as minute pores only. Wax glands present ..
............................................................. 4
SIPH present as tubes or large pores on broad hairy cones. Wax glands
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
4 R IV+V 0.5 x HT II or less. CAUDA bluntly triangular. Alata with
dark transverse bars on abdomen, usually with a once-branched media
and secondary rhinaria on ANT III only (sometimes 1-2 distally on IV)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mindarus abietinus
 Uapaca 509

- R IV+V equal to or longer than HT II. CAUDA not bluntly triangular.

Alata (sexupara) with no dorsal abdominal markings, an unbranched
media and secondary rhinaria on ANT III-V or III-VI . . . . . . . . . . . 5
5 R IV+V more than 0.16mm. Alate sexupara with BL 2.4-3.0mm, and
ABD TERG 8 extended as anocaudal process (as in Fig. 68A) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prociphilus taxus
R IV+V less than 0.14mm. Alate sexupara with BL 1.1-1.9 mm and
without an anocaudal process . . . . . . . . . . . . . . . . . . . . . Prociphilus pini
6 SIPH long, tubular, slightly swollen on distal half with a subapical, con-
stricted zone of polygonal reticulation. CAUDA long, finger-like, about
0.5 x SIPH. Hairs on body and appendages very short and blunt . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia patriciae
SIPH as large pores on broad hairy cones. CAUDA rounded. Hairs on
body and appendages mainly long and pointed . . . . . . . . . . . . . . . . . . . 7
7 Hind tibiae pale except distally. SIPH cones pale . . . Cinara pilicornis
Hind tibiae mainly dark. SIPH cones strongly pigmented . . . . . . . . . 8
8 BL 3.8-4.2mm. Hairs on ABD TERG 2-7 (probably) long. Alata with
secondary rhinaria distributed ANT III 4-5, IV 2, V 0-1 . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara taiwana*
- BL 2.9-3.5 mm. Hairs on ABD TERG 2-7 very short and blunt. Alata
with secondary rhinaria distributed ANT III 5-8, IV 1-2, V 2 . . . . . . .
.................................................. Cinara tsugae

TURPINIA Staphyleaceae

Turpinia pomifera Indomegoura indica

UAPACA Euphorbiaceae

Uapaca guineensis Aphis gossypii

510 Ulmus

ULMUS Elms Ulmaceae

In all, 68 aphid species are recorded from elms, which are primary hosts for
most of the Eriosomatini (44 species). For gall formers, Eriosomatini show
remarkably little evidence of host specificity, the recorded host range reflect-
ing geographical distribution rather than host affinities, with the more widely-
distributed species such as Eriosoma ulmi recorded from Ulmus species native
to different parts of the world. Elm-feeding aphids are therefore all included
in a single key as far as possible. However, at the present time it is not possible
to key together all species of the large genus Eriosoma and its close relatives
(Georgiaphis, Schizoneurella), so these are taken out of the main key and
included in separate keys under particular host species, within the host plant
list that follows the main key.

Main key to elm aphids

1 CAUDA knobbed, anal plate bilobed. SIPH as short truncate cones
without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- CAUDA if present broadly rounded, tongue-like or finger-like. Anal plate
if present entire or with a median cleft but not distinctly bilobed. SIPH
if present not in form of truncate hairless cones . . . . . . . . . . . . . . . . 14
2 Colonies include apterous viviparae, with dark markings and long dorsal
hairs, at least some of which have furcate apices . . . . . . . . . . . . . . . . . 3
- All adult viviparae alate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3 Aptera with longest hairs on ANT III shorter than basal diameter of
segment, and with spinopleural sclerites of ABD TERG 1-6 mutually free
and well separated from marginal sclerites . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Indochaitophorus furcatus
- Aptera with longest hairs on ANT III longer than basal diameter of
segment, and ABD TERG 1-6 more-or-less fused between segments
............................................................. 4
4 Aptera with longest hairs on ANT III up to 2.5 x basal diameter of
segment. Dorsum with paler spinal region and with pale areas around
SIPH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinochaitophorus mordvilkoi
- Aptera with longest hairs on ANT III more than 3 x basal diameter of
segment. Dorsum wholly dark except for a small median patch, with SIPH
incorporated into pigmented area . . . . . . . . . . . Sinochaitophorus maoi
5 ANT PT/BASE about 0.5 or less. Tibiae wholly dark and rather thick
with long hairs. Secondary rhinaria transversely oval. Spinal hairs on
ABD TERG 1 and 2 (and pleural hairs also on ABD TERG 2) usually
arising from low tubercles . . . . . . . . . . . . . . . . . . . Chromocallis nirecola
 Ulmus 511

- ANT PT/BASE 0.6 or more. Tibiae thin and mainly pale with mostly
short hairs. Secondary rhinaria very narrow, slit-like. Spinal hairs on ABD
TERG 1 and 2 on prominent finger-like or conical tubercles (pleural hairs
absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Media of forewing once-branched, with a patch of fuscous distally
between the branches, and another at the distal end of Cu lb (Fig. 116C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis (Sappocallis) ulmicola
- Forewing with media twice-branched, and with or without a different
pattern of markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Head with 3 pairs of dorsal tubercles, bearing apical hairs; the most
posterior pair longest, finger-like (Fig. 116A) . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis ulmiparvifoliae
- Head without paired dorsal tubercles or with only one very small pair
............................................................. 8
8 Forewing with media and CU1b thickly and rather evenly bordered with
fuscous basally as well as distally, and Cu la also often partially bordered.
Abdomen robust, with more-or-less extensive dark dorsal markings. SIPH
dark, with a dark basal sclerite . . . . . . . . . . . . . . . . . . Tinocallis platani
- Forewing with veins mostly not bordered or with fuscous restricted to
branches of media distally, or to spots or patches at distal ends of veins
............................................................. 9
9 Pronotum with 1 (posterior) or 2 pairs of finger-like spinal tubercles
(e.g. Fig. 116B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Pronotum without spinal tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10 Head and pronotum wholly black (dark brown in mounted specimens),
and dorsal abdomen without dark markings . . . . . . . . . . . . . . . . . . . . 11
- Head and pronotum, or at least pronotum, pale or dusky or with restricted
dark markings. ABD TERG 3-5 (or 3-7) with dark sclerites at bases of
spinal hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Forewings with distal branches of media bordered with fuscous and with
more-or-less extensive fuscous patches at distal ends of Cula and Cu lb
(Fig. 51B). ANT III with 14-24 secondary rhinaria. R IV+V with 6-10
accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis takachihoensis
- Forewings without fuscous markings (or, at least, none discernible in the
mounted specimens available). ANT III with 10-12 secondary rhinaria.
R IV+V with 4-6 accesory hairs . . . . . . . . . . . . Tinocallis sapporoensis
12 Distal branches of media of forewing bordered with fuscous, and a
patch of fuscous also on the hind margin at the end of Cu1b (Fig. 116D).
R IV+V 0.76-0.90 x HT II. ANT III with 6-17 secondary rhinaria
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis saltans
512 Ulmus

Fig. 116. A, head of Tinocallis ulmiparvifoliae; B, prothorax of T. saltans; C, forewing of T. ulmicola;

D, forewing of T. saltans.

- Forewing veins not bordered with fuscous, and hind margin without a
patch of fuscous at end of Cu lb . R IV+V 0.90-1.03 x HT II. ANT III
with 11-28 secondary rhinaria . . . . . . . . . . . . . . . . . . . Tinocallis nevskyi
13 Head with a single narrow dark dorsal mesial stripe. ANT III with 14-22
secondary rhinaria. Spinal tubercles on ABD TERG 1 and 2 shorter than
their basal widths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis zelkowae
- Head with variable pigmentation; sometimes with paired longitudinal
stripes but never with a single narrow mesial stripe. ANT III with 5-12
secondary rhinaria. Spinal tubercles on ABD TERG 1 and 2 large and
prominent, usually larger than their basal widths, sometimes united at
bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis ulmifolii
14 Body of aptera broadly pear-shaped, broadest anteriorly, without distinct
head and prothorax. Oviparous. (Alata undescribed) . . . . . . . . . . . . . . .
............................................... Olegia ulmifoliae
- Body of aptera not broadest anteriorly, head and prothorax distinct
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Rostrum much longer than body. SIPH as large pores on broad pigmented
hairy cones. Abdomen with a mesial row of 6 dark patches ventrally
............................................. Stomaphis ulmicola
- Rostrum much shorter than body. SIPH variously shaped . . . . . . . . 16
 Ulmus 513

16 SIPH tubular, much longer than basal width. ANT PT/BASE more than
1. Wax glands absent . . . . . . go to key to polyphagous aphids, p. 532
- SIPH absent, ring- or pore-like, or raised on small shallow cones. ANT
PT/BASE less than 0.6. Wax glands often present . . . . . . . . . . . . . . 17
17 Apterae in brown microrrhizal cysts on roots, with distal hairs on tibiae
thick and spine-like or dagger-like, and tarsi 1-segmented. SIPH absent
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mimeuria ulmiphila
- Apterae in galls or leaf nests, except for one species (Eriosoma rileyi)
forming free-living colonies on bark; tibial hairs normal, tarsi 1- or
2-segmented. SIPH present as pores, sometimes raised on small shallow
cones with surrounding hairs, or absent . . . . . . . . . . . . . . . . . . . . . . . . 18
18 (All remaining couplets refer only to alatae produced in or emerging from
galls or leaf nests.) BL more than 4mm. Hindwing with 2 oblique veins
arising very close together . . . . . . . . . . . . . . . . . . . . . . Prociphilus oriens
- BL less than 4 mm. Hindwing if with 2 oblique veins then their bases are
separated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19 Hindwing with 2 oblique veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
- Hindwing with 1 oblique vein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
20 SIPH present as fairly large, often raised pores on ABD TERG 5, with
partially sclerotized rims and surrounding hairs. R IV+V with 4 or more
accessory hairs. ANT III elongate (Fig. 118), usually more than 0.8 x
head width across (and including) eyes (except in Eriosoma mimicum)
............................... go to keys to species of Eriosoma,
Georgiaphis and Schizoneurella under individual Ulmus spp.
- SIPH absent or present as small or large pores on ABD TERG 6, with
wholly sclerotized rims and no clearly associated hairs. R IV+V with 2-3
accessory hairs. ANT III less than 0.8 x head width across eyes (e.g.
Figs 117A-F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21 Dorsal abdomen with dark intersegmental transverse bands, broadest in
midline. ANT BASE VI much shorter than R IV+V. Secondary rhinaria
distributed ANT III 12-18, IV 2-5, V 2-4, VI 0-3 (Fig. 117A) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphidounguis mali
- Dorsal abdomen without transverse bands. ANT BASE VI much longer
than R IV+V. Secondary rhinaria distributed ANT III 9-24, IV 2-7,
V 4-12, VI 4-10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22 Embryos without mouthparts (sexupara) . . . . Kaltenbachiella japonica
- Embryos with mouthparts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23 SIPH absent. Secondary rhinaria distributed ANT III 17-25, IV 4-7,
V 5-8, VI 7-10 (Fig. 117B) . . . . . . . . . . . . . . . . . Kaltenbachiella pallida
514 Ulmus

Fig. 117. Antennae of alatae from galls of A, Aphidounguis mali, B, Kaltenbachiella pallida,
C, K. spinosa, D, Colopha compressa, E, Tetraneura ulmi, F, T. nigriabdominalis; G, SIPH of
T. yezoensis, lateral and dorsal views; H, hind claws of embryo of T. javensis (seen through cuticle of alata).

- SIPH usually present, but if sometimes absent then ANT III has fewer
than 17 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24 Secondary rhinaria distributed ANT III 17-25, IV 5-9, V 7-9, VI 6-10
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kaltenbachiella ulmifusa
- Secondary rhinaria distributed ANT III 9-17, IV 2-6, V 4-12, VI 4-10
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25 Media of forewing unbranched. Secondary rhinaria distributed ANT III
11-17, IV 4-6, V 8-12, VI 6-10 (Fig. 117C) . . . . . . . . . . . . . . . . . . . . . . .
.......................................... Kaltenbachiella spinosa
- Media of forewing usually once-branched. Secondary rhinaria distributed
ANT III 9-12, IV 2-4, V 4-6, VI 4-6 . . . . . . Kaltenbachiella nirecola
 Ulmus 515

26 ANT IV similar in length to ANT V (in 6-segmented antennae, Fig. 117D;

if antenna 5-segmented then 'ANT III' more than 3 x 'ANT IV). Media
of forewing once-branched or unbranched . . . . . . . . . . . . . . . . . . . . . . 27
- ANT IV much shorter than V (e.g. Figs 117E-F; if antenna 5-segmented
then 'ANT III' less than 2 x 'ANT IV). Media of forewing always
unbranched ................................................. 29
27 R IV+V 0.10-0.11 mm long, 0.8-0.9 x HT II. Media of forewing once-
branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colopha compressa
- R IV+V 0.05-0.07 mm, 0.6-0.75 x HT II. Media of forewing once-
branched or unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28 Media of forewing once-branched . . . . . . . . . . . . . . . . Colopha ulmicola
- Media of forewing unbranched . . . . . . . . . . . . . . . . . . Colopha graminis
29 Embryos with hind tarsal claws less than 0.05 mm long . . . . . . . . . . 30
- Embryos with enlarged hind tarsal claws at least one of which is more
than 0.06 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
30 ANT II with 9-19 hairs. R IV+V with 10-14 accessory hairs. SIPH
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetraneura radicicola/persicina
- ANT II with 2-7 hairs. R IV+V with 5-10 accessory hairs. SIPH present
or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31 SIPH always present, with dark rims (Fig. 117G). Frontal hairs long and
fine, maximally 60-80 µm long . . . . . . . . . . . . . . . . Tetraneura yezoensis
- SIPH absent or if present then with pale rims. Frontal hairs 10-35 µm
long ........................................................ 32
32 ANT V 1.0-1.25 x ANT VI (including PT). R IV+V 0.085-0.11 mm long,
about 0.6 x HT II, with 5-7 accessory hairs. SIPH and abdominal wax
glands absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetraneura caerulescens
- ANT V 1.5-2.0 x ANT VI (Fig. 117E). R IV+V 0.11-0.15mm long,
0.7-0.9 x HT II, with 6-10 accessory hairs. SIPH present or absent, if
present then there are usually also ring-like wax glands . . . . . . . . . . . . .
.......................................... Tetraneura ulmi group
33 CAUDA with 4 hairs. Embryos with only one hind tarsal claw markedly
enlarged, the shorter one less than 0.05 mm long (Fig. 117H) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetraneura javensis
- CAUDA with 2, or rarely, 3 hairs. Embryos with both hind tarsal claws
enlarged, sometimes unequally, but both more than 0.06 mm long ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
34 R IV+V 0.10-0.13 mm long, with 6-18 accessory hairs . . . . . . . . . . . 35
- R IV+V 0.08-0.105 mm long, with 2-6 accessory hairs. (Secondary
rhinaria distributed ANT III 7-19, IV 2-6, V 5-12) . . . . . . . . . . . . . 36
516 Ulmus

35 R IV+V with 14-18 accessory hairs. Secondary rhinaria distributed ANT

III 20-32, IV 3-10, V 8-15 . . . . . . . . . . . . . . . . . Tetraneura polychaeta
- R IV+V with 6-8 accessory hairs. Secondary rhinaria distributed ANT III
10-14, IV 2-3, V 9-10 . . . . . . . . . . . . . . . . . . . . Tetraneura asymmachia
36 Tarsi of embryos without spicules. ABD TERG 1-6 of embryos each with
6-12 spinopleural hairs between the stout lateral hairs . . . . . . . . . . . 37
- Tarsi of embryos minutely spiculose. ABD TERG 1-6 of embryos each
with 4-6 spinopleural hairs between the stout lateral hairs . . . . . . . . 38
37 R IV+V 0.09-0.095 mm long, 0.6-0.65 x HT II and bearing 6 acces-
sory hairs. ANT V about 2.9 x ANT VI (incl. PT). ABD TERG 1-6 of
embryos each with 10-12 fine, pointed spinopleural hairs between the
stout lateral hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tetraneura chinensis
- R IV+V 0.075-0.09 mm long, with 2-4 accessory hairs. ANT V 1.7-2.8 x
ANT VI (Fig. 117F). ABD TERG 1-6 of embryos each with 6-10 spino-
pleural hairs between the laterals . . . . . . . Tetraneura nigriabdominalis
38 R IV+V 0.08-0.09 mm long with 4 accessory hairs. Secondary rhinaria
distributed ANT III 7-10, IV 2-5, V 6-9. ABD TERG 1-6 of embryos
each with 4 spinopleural hairs + 2 stout lateral hairs. Hind tarsi of
embryos distinctly spiculose . . . . . . . . . . . . . . . . . . . . . . Tetraneura sorini
- R IV+V 0.09-0.105 mm long with 5-6 accessory hairs. Secondary rhinaria
distributed ANT III 12-18, IV 3-6, V 6-12. ABD TERG 1-6 of embryos
each with usually 6 fine spinopleural hairs + 2 stout lateral hairs. Hind
tarsi of embryos very slightly spiculose . . . . . . . . Tetraneura fusiformis

Host Plant List (including keys to Eriosoma, etc.)

Ulmus alata Eriosoma lanigerum group
Georgiaphis ulmi
Tinocallis ulmifolii

Key to Eriosomatini spp. (alatae from galls)

ANT III with secondary rhinaria rather widely and irregularly spaced,
and not usually extending half-way around segment (as in Fig. 118A).
Embryos without mouthparts . . . . . . . . . . . . . . . . . . . . Georgiaphis ulmi
ANT III with secondary rhinaria closely and regularly spaced and often
extending more than half-way around segment . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanigerum group

U. americana Colopha compressa, graminis,

(American White Elm) ulmicola
Eriosoma americanum, ?crataegi,
grossulariae, lanigerum group,
 Ulmus 517

mimicum, pyricola, rileyi, ulmi

Georgiaphis ?gillettei, [maxsoni]
Macrosiphum euphorbiae
Tinocallis platani, ulmifolii

Key to Eriosoma and Georgiaphis on Ulmus americana

1 Aphids with apterae in free-living colonies on bark . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma rileyi
Aphids forming galls (remaining couplets refer only to alatae produced
in galls) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 ANT III with secondary rhinaria rather widely and irregularly spaced
and not usually extending more than half-way around segment (Fig. 118A).
Embryos without mouthparts (no coiled stylets visible in maternal
abdomen) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Georgiaphis gillettei
ANT III with secondary rhinaria closely or at least regularly spaced and
often extending more than half-way around segment (e.g. Fig.118C).
Embryos with mouthparts (coiled stylets visible) . . . . . . . . . . . . . . . . . . 3
3 ANT III less than 0.8 x head width across (and including) eyes. R IV+V
only 0.07-0.10 mm long . . . . . . . . . . . . . . . . . . . . . . . Eriosoma mimicum
- ANT III more than 0.8 x head width across eyes. R IV+V 0.13-0.26 mm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 ANT V with secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
ANT V without secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
5 ANT III a little shorter than ANT IV+V+VI (Fig. 118C) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma americanum
- ANT III as long as or longer than ANT IV+V+VI . . . . . . . . . . . . . . . 6
6 ANT III 0.50-0.61 mm, ANT V 0.16-0.20 mm and ANT VI (incl. PT)
0.14-0.15 mm. Secondary rhinaria distributed III 27-34, IV 5-8, V 7-11,
VI 1-3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma crataegi
- ANT III 0.35-0.52 mm, ANT V 0.09-0.16 mm and ANT VI (incl. PT)
0.07-0.10 mm. Secondary rhinaria distributed III 20-30, IV 4-7, V 4-7,
VI 0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 CAUDA with 2 hairs. ANT V slightly longer than ANT IV. First tarsal
segments of hind legs with 3 hairs (incl. sense peg) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma lanigerum group
- CAUDA with 3-10 hairs. ANT V not longer than or slightly shorter than
IV. First tarsal segments of hind legs with 2 hairs (no sense peg) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma pyricola
8 ANT PT long and thin; PT/BASE 0.46-0.67 (Fig. 118D). Many hairs on
518 Ulmus

Fig. 118. Antennae of alatae from galls of A, Georgiaphis gillettei, B, Schizoneurella indica, C, Eriosoma
americanum, D, E. grossulariae (ANT VI only), E, E. ulmi (ANT VI only), F, E. moriokense, G, E. harunire.

ABD TERG 1-5 standing on small scleroites. HT II with 22-25 hairs

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma grossulariae
- ANT PT shorter and thicker (Fig. 118E); PT/BASE 0.29-0.49. Few hairs
on ABD TERG 1-5 with basal scleroites. HT II with 15-22 hairs . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma ulmi

U. campestris = U. procera
U. canescens Aphis spiraecola
Toxoptera aurantii
U. carpinifolia Colopha compressa
(Smooth-Leaved Elm) Eriosoma anncharlotteae, flavum,
 Ulmus 519

grossulariae, lanuginosum,
patchiae, pyricola, ulmi
Tetraneura caerulescens,
nigriabdominalis, ulmi
Tinocallis platani
(For separation of Eriosoma spp. use key under Ulmus procera, p. 532)
U. davidiana Chromocallis nirecola
Tinocallis ulmicola
Tetraneura nigriabdominalis
U. davidiana var. japonica = U. japonica
U. densa Aphis craccivora, [sogdiana
Nevsky, 1929b]
Eriosoma phaenax
Tetraneura ulmi
U. effusa Colopha compressa
(Russian White Elm) Eriosoma ulmi
Tetraneura ulmi
Tinocallis platani
U. foliacea = U. carpinifolia
U. fulva = U. rubra
U. glabra Aulacorthum solani
(Wych Elm) Colopha compressa
Eriosoma flavum, grossulariae,
lanuginosum, patchiae,
phaenax, ulmi, ?ulmosedens
Kaltenbachiella japonica, pallida
Tetraneura caerulescens, ulmi
Tinocallis nevskyi, platani,
takachihoensis
Toxoptera aurantii
(For separation of Eriosoma spp. use key under Ulmus procera, p. 532)
U. japonica Chromocallis nirecola
(Harunire, Japanese Elm) Colopha nirecola
Eriosoma alabastrum, auratum,
eligulatum, harunire, japonica,
longicornutum, mediocornutum,
moriokense, ulmi, yangi ssp.
parasiticum
Kaltenbachiella japonica, nirecola,
spinosa
Olegia ulmifoliae
Prociphilus kuwanai, oriens
Sinochaitophorus mordvilkoi
520 Ulmus

Stomaphis ulmicola
Tetraneura fusiformis,
nigriabdominalis, radicicola,
sorini, [triangula], ulmi,
yezoensis
Tinocallis saltans, takachihoensis,
ulmicola, zelkowae

Key to alatae of Eriosoma spp. from galls on Ulmus japonica

1 ANT V with 5-7, and ANT VI with 1-2, secondary rhinaria . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alabastrum *
- ANT V with 0-3, and ANT VI without secondary rhinaria . . . . . . . . 2
2 ANT III shorter than IV+V+VI added together, and bearing 10-20 secon-
dary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- ANT III longer than IV+V+VI added together (e.g. Figs 118F, G), and
bearing 22-46 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Total antennal length less than 0.6mm, with ANT III 0.19-0.23 mm.
Secondary rhinaria distributed III 10-13, IV 1-2, V 0, VI 0 . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . yangi ssp. parasiticum
Total antennal length greater than 0.7 mm, with ANT III 0.27-0.34 mm.
Secondary rhinaria distributed III 13-20, IV 1-5, V 0-1, VI 0 . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . japonicum
4 ANT V and VI both very short, each less than 0.5 x ANT IV (Fig. 118F).
(Secondary rhinaria distributed III 33-41, IV 5-9, V 0, VI 0) . . . . . . .
.................................................... moriokense
ANT V and VI both similar in length to ANT IV, or only VI somewhat
shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Total antennal length 1.4-1.8 mm. ANT PT 0.06-0.08 mm, with ANT
PT/BASE more than 0.5. (Secondary rhinaria distributed III 36-46,
IV 6-8, V 0, VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . longicornutum
- Total antennal length 0.70-1.32 mm. ANT PT 0.02-0.06 mm, with ANT
PT/BASE less than 0.5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 ANT V with 0-3 secondary rhinaria (usually at least one on one side).
Secondary rhinaria all very narrow and thin, so that antenna not serrate
in profile (Fig. 118G); distributed III 24-30, IV 3-5, V 0-3, VI 0 . . . .
....................................................... harunire
ANT V never with secondary rhinaria. Secondary rhinaria on ANT
III-IV broad and thick, giving profile of antenna a serrate appearance
............................................................. 7
7 ANT PT 0.02-0.03 mm. Secondary rhinaria distributed III 22-30, IV 3-6,
V 0, VI 0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . auratum
 Ulmus 521

- ANT PT 0.04-0.06 mm (Fig. 118E). Secondary rhinaria distributed III

28-41, IV 6-9, V 0, VI 0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ulmi

U. laciniata Eriosoma laciniatae,

longicornutum
Kaltenbachiella japonica
Tetraneura sorini, yezoensis
Tinocallis sapporoensis,
takachihoensis

Key to Eriosoma spp.

Primary rhinarium on ANT V without a ciliated rim. Secondary rhinaria
distributed III 24-29, IV 4-6, V 0, VI 0. ANT PT/BASE less than 0.4
...................................................... laciniatae
Primary rhinarium on ANT V ciliated. Secondary rhinaria distributed
III 36-46, IV 6-8, V 0, VI 0. ANT PT/BASE more than 0.4 . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . longicornutum

U. laevigata Eriosoma kashmiricum

Indiochaitophorus furcatus
Schizoneurella indica
Tetraneura polychaeta
Tinocallis saltans

Key to separate alatae of Eriosoma and Schizoneurella from galls on Ulmus

laevigata
- R IV+V 0.11-0.13 mm, with 4-6 accessory hairs. ANT V and VI with
rhinarial complexes (formed by fusion of primary and secondary rhinaria)
of very irregular shape, and not conspicuously ciliated (Fig. 118B).
Secondary rhinaria distributed ANT III 19-29, IV 4-8, V 0-5, VI 0-2
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoneurella indica
- R IV+V 0.15-0.18 mm, with 18-20 accessory hairs. ANT V and VI
spinulose with primary rhinaria normal, round, ciliated. Secondary
rhinaria distributed III 22-32, IV 4-7, V 0, VI 0 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eriosoma kashmiricum

U. laevis Colopha compressa

Tinocallis platani
U. macrocarpa Kaltenbachiella japonica
Tinocallis takachihoensis
U. mandshurica Aphidounguis mali
Tetraneura nigriabdominalis
[Tinocallis coreanus]
522 Ulmus

U. montana = U. glabra
U. parvifolia Aphidounguis mali
(Akinire, Chinese Elm) Eriosoma yangi
Tetraneura nigriabdominalis,
sorini, yezoensis
Tinocallis ulmiparvifoliae,
zelkowae
U. pedunculata Colopha compressa
U. pinnatoramosa Eriosoma ulmi
Tetraneura ulmi
Tinocallis platani
U. procera Aphis craccivora, [sogdiana
(English Elm) Nevsky, 1929b]
Chromocallis nirecola
Colopha nirecola
Eriosoma anncharlotteae, flavum,
grossulariae, japonicum,
lanuginosum, patchiae,
phaenax, pyricola, ulmi,
ulmosedens
Kaltenbachiella japonica, pallida
Mimeuria ulmiphila
Tetraneura africana, caerulescens,
nigriabdominalis, ulmi,
yezoensis
Tinocallis nevskyi, platani,
saltans, ulmicola, ulmifolii,
ulmiparvifoliae

Key to alatae of Eriosoma spp. from galls on Ulmus procera (including also
species on U. carpinifolia and U. glabra)
1 Antenna with only 14-28 secondary rhinaria in total (distributed III
12-20, IV 2-5, V 0-4, VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Antenna with at least 29 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . 3
2 ANT III shorter than ANT IV+V+VI. Secondary rhinaria on III mostly
not extending more than half-way around segment . . . . . . . japonicum
ANT III longer than ANT IV+V+VI. Secondary rhinaria on III mostly
extending more than half-way around segment . . . . . . . . . . . . . . flavum
3 ANT V without secondary rhinaria. (Secondary rhinaria distributed III
26-46, IV 3-8, V 0, VI 0) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
ANT V with secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
 Ulmus 523

4 Total antennal length less than 1 mm, with ANT III less than 0.6 mm.
HT 0.15-0.18 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . anncharlotteae
Total antennal length more than 1 mm, with ANT III more than 0.6 mm.
HT 0.17-0.21 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 ANT PT long and thin; PT/BASE 0.46-0.67 (Fig. 118D). Many hairs on
ABD TERG 1-5 standing on small scleroites. HT II with 22-25 hairs
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . grossulariae
- ANT PT shorter and thicker; PT/BASE 0.29-0.49 (Fig. 118E). Few hairs
on ABD TERG 1-5 with basal scleroites. HT II with 15-22 hairs . . . .
.......................................................... ulmi
6 ANT V longer than IV. R IV+V 1.20-1.35 x HT II, and bearing 18-23
accessory hairs. Apices of tibiae and first tarsal segments strongly spiculose.
(Secondary rhinaria distributed III 18-35, IV 2-7, V 1-7, VI 0) .... 7
- ANT V shorter than or as short as IV. R IV+V 0.8-1.2 x HT II and
bearing 8-14 accessory hairs. Apices of tibiae and first tarsal segments
with only a few minute spinules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7 Alatae are sexuparae, having embryos without mouthparts . . . . . . . . . .
.................................................... ulmosedens
Alatae have embryos with mouthparts (coiled stylets visible through
maternal cuticle) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . patchiae
8 Secondary rhinaria distributed ANT III 33-48, IV 10-14, V 10-12,
VI 2-4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . phaenax
- Secondary rhinaria distributed ANT III 21-35, IV 7-11, V 5-16, VI 0-1
............................................................. 9
9 BL 2.1-3.1 mm. R IV+V 0.19-0.23 mm. CAUDA with only 2 or rarely
3 hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lanuginosum
- BL 1.3-2.4 mm. R IV+V 0.10-0.19 mm. CAUDA with 3-10 hairs . . . .
....................................................... pyricola

U. propinqua = U. japonica
U. pubescens = U. fulva
U. pumila Chromocallis nirecola, [pumili
(Asiatic Dwarf Elm) Zhang, 1982 (in Zhang and
Zhong, 1982c)], [similinirecola
Zhang, 1982 (in Zhang and
Zhong, 1982e)],
Eriosoma dilanigiosum,
eligulatum, mediocornutum,
ulmipumilae
Sinochaitophorus maoi
Tetraneura asymmachia, chinensis,
nigriabdominalis
Tinocallis saltans
524 Ulmus

Key to alatae of Eriosoma spp. from galls on Ulmus pumila

1 ANT V, or ANT V and VI, with secondary rhinaria . . . . . . . . . . . . . 2
ANT V and VI both without secondary rhinaria . . . . . . . . . . . . . . . . . 3
2 ANT V with 5-7 and ANT VI with 1-2 secondary rhinaria . . . . . . . . .
.................................................. dilanigiosum *
- ANT V with 2-3 and ANT VI with 0 secondary rhinaria . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ulmipumilae*
3 ANT PT about 0.03 mm. Secondary rhinaria distributed III 22-26, IV 3-5
.................................................... eligulatum*
- ANT PT 0.04mm or more. Secondary rhinaria distributed III 28-36,
IV 6-8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . mediocornutum *
(or try key to Eriosoma spp. on U. japonica, p. 520)

U. racemosa = U. thomasii
U. rubra Colopha graminis, ulmicola
(Slippery Elm) Eriosoma ?crataegi, mimicum,
rileyi
Georgiaphis gillettei
Kaltenbachiella ulmifusa
(For separation of Eriosoma and Georgiaphis spp. use key under Ulmus
americana, p. 517)
U. scabra = U. glabra
U. suberosa = U. procera
U. thomasii Colopha compressa, ulmicola
Eriosoma ulmi
Tinocallis platani
U. uyematsui Eriosoma lishanense
Kaltenbachiella glabra
U. wallichiana Eriosoma kashmiricum, phaenax
Indiochaitophorus furcatus
Tetraneura javensis
Tinocallis saltans

Key to alatae of Eriosoma spp. from galls on Ulmus wallichiana

- Secondary rhinaria distributed ANT III 33-48, IV 10-14, V 10-12, VI 2-4.
R IV+V with 12-14 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . phaenax
- Secondary rhinaria distributed ANT III 22-32, IV 4-7, V 0, VI 0. R IV+V
with 18-20 accessory hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . kashmiricum

Ulmus spp. Aphis gossypii

Brachycaudus helichrysi
 Vernonia 525

Myzus ornatus, persicae

[Phorodon humulifoliae Tseng
and Tao, 1938]
Tetraneura [aequiunguis],
[changaica], persicina, triangula

UMBELLULARIA Lauraceae

Umbellularia californica Aulacorlhum circumflexum

Euthoracaphis umbellulariae
Myzus persicae

Key to species on Umbellularia (apterous viviparae)

Body aleyrodiform; subcircular, heavily sclerotized, dark, with reduced
legs and antennae, and bearing numerous long fine dorsal and marginal
hairs (Fig. 39A) . . . . . . . . . . . . . . . . . . . . . . . Euthoracaphis umbellulariae
Body of normal aphid form . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . go to key to polyphagous aphids, p. 532

UVARIA Annonaceae

Uvaria sp. Greenideoida ceyloniae ssp.

bhalukpongensis

VEPRIS Rutaceae

Vepris lanceolata Toxoptera citricidus

V. undulata Toxoptera citricidus

VERNONIA Compositae

Several African Sitobion spp. occur on herbaceous Vernonia spp.

526 Virgilia

Vernonia arborea Uroleucon compositae

V. leucocalyx Uroleucon compositae

VIRGILIA Leguminosae

Virgilia oroboides Aphis craccivora

VITEX Verbenaceae

Vitex agnus-castus Aphis viticis

Myzus persicae
V. cannabifolia Aphis gossypii
V. trifolia Aphis gossypii

Key to aphids on Vitex

1 Antennal tubercles of aptera well developed, with inner faces convergent
in dorsal view. SIPH pale, slightly swollen on distal half (Fig. 121H)
................................................. Myzus persicae
Antennal tubercles not developed. SIPH mainly dark (except in very
small, aestivating specimens), and tapering . . . . . . . . . . . . . . . . . . . . . . 2
2 R IV acute, with sides slightly concave in dorsal view; width at base at
least 2.1 x distal width at articulation with R V. SIPH usually pale at
base, darkening towards apex. Alata with 3-16 (usually 6-12) secondary
rhinaria on ANT III and 0-2 on ANT IV . . . . . . . . . . . . . Aphis viticis
R IV+V normal, with sides slightly convex in ventral view; width at base
less than 1.9 x distal width at articulation with V. Alata usually with 4-8
secondary rhinaria on ANT III and 0-2 (usually 0) on IV . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii

WIDDRINGTONIA Cupressaceae

Widdringtonia juniperoides Cinara tujafilina

[Siphonatrophia cupressi]
 Zanthoxylum 527

W. schwartzii Illinoia morrisoni

W. whytei Cinara tujafilina

Key to aphids on Widdringtonia

- ANT PT/BASE more than 3. SIPH tubular, long, markedly swollen on
distal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Illinoia morrisoni
- ANT PT/BASE less than 0.5. SIPH as large pores on broad pigmented
hairy cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinara tujafilina
(or try key to aphids on Cupressus, p. 161).

XANTHOPHYLLUM Polygalaceae

Xanthophyllum stipitatum Anomalosiphum murphyi

XYLOPIA Annonaceae

Xylopia aromatica Aphis spiraecola

ZANTHOXYLUM Rutaceae

Host Plant List

Zanthoxylum bungei Aphis gossypii
Myzus persicae
Z. flavum Toxoptera aurantii
Z. monophyllum Aphis spiraecola
Z. nitidum Toxoptera odinae
Z. piperitum [Myzus xanthomelii Shinji, 1941]
Toxoptera odinae
Z. scandens Toxoptera victoriae
Z. simulans = Z. bungei
528 Zelkova

Zanthoxylum sp. Toxoptera citricidus

(Use key to polyphagous aphids, p. 532)

ZELKOVA Ulmaceae

The small but probably ancient aphid fauna of Zelkova is poorly studied. The
gall makers span 4 or 5 genera of Eriosomatini, all distinct from those on
Ulmus ('Colopha` caucasica probably does not belong in that genus), and are
probably relicts of a much larger fauna in the past. Tinocallis on Zelkova
include some that also feed on Ulmus (nirecola, ulmiparvifoliae), plus several
others that are included in the key rather tentatively as they have not been
seen by the authors. There are certainly other Tinocallis still to be described
from Zelkova. Note that Tinocallis zelkovae and T. zelkowae are separate
species.

Host Plant List

Zelkova acuminata Aphis craccivora
Byrsocryptoides zelkovae
Colopha caucasica
Z. carpinifolia Byrsocryptoides zelkovae,
zelkovaecola
Colopha caucasica
Tinocallis zelkovae
Z. crenata [Tinocallis nevskyi]
Z. davidii = Hemiptelea davidii
Z. formosana Myzus persicae
Tinocallis mushensis, viridis
Z. schneideriana Tinocallis allozelkowae, sophorae,
suzhouensis
Toxoptera aurantii
Z. serrata Colophina arma, clematis
(Keyaki) Hemipodaphis persimilis
Paracolopha morrisoni
Stomaphis yanonis
Tinocallis zelkowae

Key to aphids on Zelkova

1 CAUDA knobbed, anal plate bilobed. SIPH as short truncate cones
without hairs. (Adult viviparae all alate) . . . . . . . . . . . . . . . . . . . . . . . . 2
 Zelkova 529

Fig. 119. A, head of alata of Tinocallis viridis; dorsal abdominal markings of B, T. zelkovae and
C, T. zelkowae.

- CAUDA rounded, anal plate entire. SIPH absent or if present then not
in form of short truncate cones without hairs. (Adult viviparae apterous
and alate) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2 Head with all 4 pairs of dorsal cephalic hairs borne on conical tubercles
(Fig. 119A). (ANT III with 15-26 secondary rhinaria) . . . . . . . . . . . . . . .
................................................ Tinocallis viridis
- Head without dorsal tubercles, or with only one very small pair . . . . . 3
3 ANT III about as long as IV+V+VI together and bearing about 40
secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis mushensis*
- ANT III 0.5-0.7 x ANT IV+V+VI together and bearing 4-25 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Forewing with media and Cu lb thickly bordered with fuscous, especially
distally, and Cula with a distal fuscous patch. Dorsal abdomen with
extensive dark markings (Fig. 119B). ANT III with 4-13 secondary
rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis zelkovae
- Forewing veins not thickly bordered with fuscous; sometimes with small
spots at distal ends. Dorsal abdomen without extensive dark markings;
sometimes with spots at bases of hairs (e.g. Fig. 119C). ANT III with
10-25 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Thorax without spinal tubercles, and ABD TERG 1-2 with short conical
spinal tubercles, shorter than their basal widths (Fig. 119C). (Head with
a single, narrow medial longitudinal dark stripe; ANT III with 13-22
secondary rhinaria) . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis zelkowae
- Thorax with spinal tubercles, and spinal tubercles on ABD TERG 1-2
finger-like, longer than their basal widths . . . . . . . . . . . . . . . . . . . . . . . 6
6 Spinal hairs on ABD TERG 3-7 without pigmented bases. ANT III with
20-25 secondary rhinaria . . . . . . . . . . . . . . . . . . . Tinocallis suzhouensis*
- Bases of spinal hairs on ABD TERG 3-7 encircled by pigment. ANT III
with 10-18 secondary rhinaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
530 Zelkova

7 BL less than 2 mm. ANT III about as long as ANT IV+V together and
bearing 10-18 secondary rhinaria extending over 0.7 or more of length
............................................ Tinocallis sophorae*
- BL more than 2 mm. ANT III much shorter than ANT IV+V together and
bearing 12-13 secondary rhinaria confined to basal half ............
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tinocallis allozelkowae*
8 BL more than 4 mm. Rostrum much longer than body. Abdomen with a
medial ventral row of 5 dark patches . . . . . . . . . . . . Stomaphis yanonis
- BL less than 3 mm. Rostrum much shorter than body. Abdomen without
ventral dark markings except posteriorly . . . . . . . . . . . . . . . . . . . . . . . . 9
9 (Remaining couplets refer to alatae emerging from leaf-roll or pouch
galls.) Media of forewing once- or twice-branched. Hindwing with 1 or
2 oblique veins. SIPH pores present . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
- Media of forewing unbranched. Hindwing always with only 1 oblique
vein. SIPH pores present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10 ANT III about as long as IV+V+VI together and bearing 28-43 secondary
rhinaria often nearly encircling segment, and on thickened ridges
giving a serrate profile (Fig. 120A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
- ANT III (of 6-segmented antenna) shorter or longer than IV+V+VI
together but bearing only 10-22 narrow secondary rhinaria, mostly not
encircling more than 0.7 of segment . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
11 Total length of antennal flagellum (III-VI) 1.2-1.4 mm. ANT PT
0.01-0.02 mm long, with quadrate apex (Fig. 120A). Secondary rhinaria
distributed ANT III 31-43, IV 9-14, V 10-16, VI 8-11 . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colophina clematis
- Total length of antennal flagellum 0.9-1.1 mm. ANT PT very short and
indistinct, less than 0.01 mm long, with rounded apex. Secondary rhinaria
distributed ANT III 28-37, IV 8-13, V 9-13, VI 5-8 . . . . . . . . . . . . . . .
................................................ Colophina arma
12 R IV+V 0.15-0.17 mm long, more than 1.3 x HT II. Secondary rhinaria
distributed ANT III 12-22, IV 2-4, V 2-4, VI 0-1 . . . . . . . . . . . . . . . . .
......................................... Hemipodaphis persimilis
- R IV+V 0.07-0.10 mm long, less than 0.8 x HT II. Secondary rhinaria
distributed ANT III 10-13, IV 2-3, V 2-3, VI 0 . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Colopha caucasica
13 ANT VI BASE (or V in 5-segmented antennae) with 3-7 secondary
rhinaria. ANT PT very short and broad, about 0.01 mm or less, with
rounded apex (Fig. 120B). SIPH pores absent. Secondary rhinaria dis-
tributed ANT III 7-16, IV 4-6, V 3-8, VI 3-7 . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracolopha morrisoni
 Ziziphus 531

Fig. 120. Antennae of alatae from galls of A, Colophina clematis, B, Paracolopha morrisoni,
C, Byrsocryptoides zelkovaecola, D, B. zelkovae.

- ANT VI BASE with 0-1 secondary rhinaria. ANT PT about 0.02mm

or more, with rounded or pointed apex. SIPH pores present. Secondary
rhinaria distributed ANT III 7-11, IV 2-3, V 0-4, VI 0-1 . . . . . . . . 14
14 ANT V about equal in length to (i.e. less than 1.2 x) ANT IV (Fig. 120C).
(Secondary rhinaria distributed ANT III 8-11, IV 2-3, V 0-3, VI 0-1)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Byrsocryptoides zelkovaecola
- ANT V 1.4-1.8 x longer than ANT IV (Fig. 120D). (Secondary rhinaria
distributed ANT III 7-11, IV 2-3, V 1-4, VI 0-1) . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Byrsocryptoides zelkovae

ZIZIPHUS Rhamnaceae

Ziziphus lotus Aphis gossypii

Myzus persicae
Z. spinachristi Aphis gossypii, zizyphi
(= nasturtiil)
Ziziphus sp. Cervaphis schouteniae
532 Polyphagous Aphids

Key to aphids on Ziziphus

1 Body with very long branched hair-bearing processes (as in Fig. 92A).
SIPH about 1.0-1.5 x hind tibia and bearing a subapical ring of hairs
........................................... Cervaphis schouteniae
Body without long hair-bearing processes, SIPH shorter and without hairs
............................................................. 2
2 Antennal tubercles well developed, with inner faces convergent in dorsal
view (Fig. 42I). SIPH slightly swollen on distal half (Fig. 121H) . . . . . .
................................................. Myzus persicae
Antennal tubercles undeveloped. SIPH tapering . . . . . . . . . . . . . . . . . . 2
3 SIPH mainly pale, dark only at apex. Some of ventral hairs on hind femur
long and fine, exceeding diameter of femur at base. Alata with secondary
rhinaria distributed ANT III 8-12, IV 2-4 . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis zizyphi (= nasturtii?)
SIPH wholly dark. Hind femoral hairs all rather short. Alata with 3-8
secondary rhinaria on ANT III and 0-1 on IV . . . . . . . Aphis gossypii

KEY TO POLYPHAGOUS TREE-DWELLING APHIDS

There are about 20 polyphagous aphid species that feed typically on herbs or
shrubs, but occasionally occur on trees. Apart from Longistigma caryae,
which is a bark feeder, they are most frequently found on young growth of
saplings, or on sucker growth of older trees. The key that follows is intended
to enable such species to be identified. It should be used only after the host
plant lists and tree-specific keys have been consulted, and in conjunction with
the species descriptions in Section HI of the book. It applies to apterae only,
except where otherwise stated. Photographs of slide-mounted preparations of
all these species except Aphis eugeniae and Myzus antirrhinii can be found in
B & E, 1984, pp.415-466.

1 ANT PT/BASE much less than 1. SIPH on flat dark hairy cones
(Fig. 121A). Very large aphid (BL more than 5 mm) . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Longistigma caryae
- ANT PT/BASE much more than 1, and usually more than 2. SIPH
tubular, of varying shape. BL less than 4 mm . . . . . . . . . . . . . . . . . . . . 2
2 SIPH pale at least on basal third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- SIPH wholly and uniformly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3 CAUDA long, dark and pointed, as long as or slightly longer than SIPH,
which are dark on about distal two thirds (Fig. 121B) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sinomegoura citricola
 Polyphagous Aphids 533

- CAUDA pale or dusky, much shorter than SIPH, which are pale or dark
only at apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 SIPH long, with a subapical zone of polygonal reticulation extending
over about distal 0.15 of length; CAUDA long and pale (Fig. 121C)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macrosiphum euphorbiae
- SIPH without any subapical polygonal reticulation . . . . . . . . . . . . . . . 5
5 SIPH slightly to moderately swollen over distal 0.5-0.7 of length (Figs
121D-H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- SIPH tapering from base to apex, without any trace of swelling (Figs
121I-K) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
6 SIPH markedly inflated over distal two thirds, the swollen part being
smooth and of maximum diameter more than 1.5 x the minimum dia-
meter of the stem; CAUDA triangular, usually with 5 hairs (Fig. 121D)
.................................... Rhopalosiphoninus staphyleae
- SIPH slightly to moderately inflated, the swollen part being weakly or
strongly imbricated and of maximum diameter less than 1.5 x minimum
diameter of stem; CAUDA triangular or finger-like with 6-8 hairs
(Figs121E-H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Head and ANT I and II dark, ANT III with 1-2 secondary rhinaria near
base. Legs mostly dark, and SIPH conspicuously dark-tipped (Fig. 121E)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorihum magnoliae
- Head and ANT I and II pale, ANT III without secondary rhinaria. Legs
mostly pale, SIPH not conspicuously dark-tipped . . . . . . . . . . . . . . . . 8
8 SIPH less than 0.8 x ANT III, and coarsely imbricated even on swollen
part; CAUDA triangular, about 1.5 x its basal width (Fig. 121F) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus (Sciamyzus) cymbalariae
- SIPH more than 0.9 x ANT III, and only weakly to moderately imbri-
cated. CAUDA finger-like, about 2 x its basal width . . . . . . . . . . . . . 9
9 Usually dark green in life. SIPH usually slightly dusky overall, with
maximum width of swollen part usually more than 0.11 x SIPH length
(Fig. 121G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus antirrhinii
- Usually pale green, yellow-green or straw-coloured in life (but darker
green in cold conditions). SIPH usually quite pale except at apices, with
maximum width of swollen part usually less than 0.11 x SIPH length
(Fig. 121H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus persicae
10 Dorsal abdomen with a large dark roughly horseshoe-shaped patch
(Fig. 121I) . . . . . . . . . . . . . . . . . Aulacorthum (Neomyzus) circumflexum
- Dorsal abdmen without a large horseshoe-shaped patch . . . . . . . . . . 11
11 Dorsal abdomen with an intersegmental pattern of dark ornamentation
(Fig. 121J). ANT III without any secondary rhinaria. ANT PT/BASE less
 534 Polyphagous Aphids

Fig. 121. End of abdomen of apterous vivipara of A, Pterochloroides persicae, B, Sinomegoura citricola,
C, Macrosiphum euphorbiae, D, Rhopalosiphoninus staphyleae, E, Aulacorthum magnoliae, F, Sciamyzus
cymbalariae, 6, Myzus antirrhinii, H, M. persicae, I, Aulacorthum circumflexum, J, Myzus ornatus,
K, Aulacorthum solani, L, Uroleucon compositae, M, U. ambrosiae, N, Aphis craccivora, 0, Toxoptera
odinae, P, T. aurantii, Q, T. citricidus, R, Aphis fabae group, S, A. nerii, T, A. spiraecola, U, A. gossypii.
 Polyphagous Aphids 535

than 2.5. SIPH tapering, with a shallow S-curve, and coarsely imbricated.
BL less than 2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myzus ornatus
- Dorsal abdomen without any dark markings. ANT III with a single
rhinarium near base. ANT PT/BASE more than 3.5. SIPH fairly straight,
weakly imbricated (Fig. 121K). BL usually more than 2mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacorthum solani
12 SIPH with a distal zone of polygonal reticulation . . . . . . . . . . . . . . . 13
- SIPH without any polygonal reticulation . . . . . . . . . . . . . . . . . . . . . . . 14
13 CAUDA black (Fig. 121L). Coxae dark. Tibiae mainly pale . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleucon compositae
- CAUDA pale (Fig. 121M). Coxae pale. Tibiae mainly dark . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Uroleucon ambrosiae
14 Dorsal abdomen almost wholly covered by an extensive solid black
sclerite; CAUDA black and usually bearing 4-7 hairs (Fig. 121N) ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis craccivora
- Dorsal abdomen without an extensive solid black sclerite. CAUDA if
black then usually with more hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15 Stridulatory mechanism present, consisting of a conspicuous pattern of
ridges on ventrolateral areas of abdominal sternites 5 and 6, and a row
of short peg-like hairs on the hind tibia (Fig. 122) . . . . . . . . . . . . . . . 16
- Stridulatory mechanism not present, although peg-like hairs are some-
times present along part of hind tibia . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
16 SIPH much shorter than (0.4-0.6 x) CAUDA (Fig. 121O). ANT PT/
BASE 2.5-3.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T o x o p t e r a odinae
- SIPH usually longer than (0.9-1.5 x) CAUDA (Figs 121P, Q). ANT
PT/BASE 3.5-5.0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17 Longest hairs on ANT III 12-27 µm long, 0.5-1.0 x basal diameter of
segment. Longest hairs on hind tibia up to 60 µm long, less than 0.6 x HT
II. CAUDA with 10-26 hairs (rarely more than 20). BL often less than
2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera aurantii
- Longest hairs on ANT III 29-64 µm long, 1.5-2.0 x basal diameter
of segment. Longest hairs on hind tibia 80-110 µm, 0.7-1.0 x HT II.
CAUDA with 19-54 hairs (rarely less than 25). BL usually more than
2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxoptera citricidus
18 Dorsum with variable sclerotic pigmentation, usually consisting of at least
cross-bands on pronotum, mesonotum and ABD TERG 7 and 8, and
small dark marginal sclerites (Fig. 121R); often also with small dark
sclerites scattered over other abdominal tergites. CAUDA with 12-19
hairs ....................................................... 19
536 Polyphagous Aphids

Fig. 122. Stridulatory ridges and pegs of Toxoptera aurantii.

- Pronotum, mesonotum and ABD TERG 7 and 8 without dark sclerotic

bands and marginal sclerites not deeply pigmented. CAUDA with 7-22
hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
19 Longest hair on ANT III 35-50 µm, longest hair on hind femur 60-85 µm.
R IV+V 0.85-1.05 x HT II. SIPH 0.9-1.6 x CAUDA and 0.10-0.17 x
BL. ANT PT/BASE 2.4-3.4 . . . . . . . . . . . . . . . . . . . . . . . . . Aphis fabae
- Longest hair on ANT III 15-25 µm, longest hair on hind femur 45-70 µm.
R IV+V 1.05-1.30 x HT II. SIPH 1.3-1.8 x CAUDA and 0.14-0.20 X
BL. ANT PT/BASE 2.9-3.4 . . . . . . . . . . . . . . . . . Aphis fabae solanella
20 HT I with 3 hairs (i.e. medial sense peg present, as on fore and mid-tarsi).
CAUDA black with 9-22 hairs (e.g. Fig. 121S) . . . . . . . . . . . . . . . . . . 21
- HT I with 2 hairs (no medial sense peg, unlike fore and mid-tarsi).
CAUDA pale, dusky or black with 4-12 hairs . . . . . . . . . . . . . . . . . . 22
21 ANT PT/BASE 3.5-4.8. R IV+V 1.3-1.6 x HT II. Hind tibiae black
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis nerii
- ANT PT/BASE 2.5-3.5. R IV+V 1.1-1.3 x HT II. Hind tibiae only dark
towards apices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis eugeniae
22 Femoral hairs long and fine, the longest of them being longer than the
diameter of the femur at its base. CAUDA black, usually with a distinct
constriction about one third from base, and bearing 6-12 hairs (Fig. 121T)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis spiraecola
 Polyphagous Aphids 537

- Femoral hairs all rather short, not exceeding the diameter of the femur at
its base. CAUDA pale or dark, but usually clearly paler than siphunculi,
tongue-shaped without a constriction, and bearing 4-7 hairs (Fig. 121U)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphis gossypii
 THE APHIDS

INTRODUCTION
In this part of the book we provide information on the aphids listed and keyed
in Section II, treating them in alphabetical order of genera, and of species
within genera. If a species listed in Section II is not found in the alphabetical
sequence then it is probably mentioned in the discussion of some other species;
the index to aphid species names (Section IV) will indicate where to look.
The classification of Aphidoidea follows that used in Blackman and
Eastop (1984), rather than that of Heie (1980) which elevates most of the
subfamilies of Aphididae to family level. Table 3 compares these two classi-
fications and shows the equivalent categories.
The information given comprises appearance in life, body length of one
or more morphs, host plants, distribution, life cycle including time of appear-
ance of sexual morphs where known, parasitoids, selected references to the
biology and economic importance, and chromosome number where known.
The range of body length (BL) gives an indication of size, but should be
treated warily for lesser-known species where it may be based on very few
specimens. References are selected mainly to provide a lead into the literature
on a particular species, so more recent publications are sometimes included
in preference to earlier more comprehensive or significant pieces of work.
Chromosome number is usually given as 2n (female), with a reference for
those chromosome numbers not included in published lists (e.g. Kuznetsova
and Shaposhnikov, 1973; Blackman, 1980); previously unpublished chromo-
some numbers are indicated by an asterisk*.

538
 Acanthochermes 539

Table 3. Classification of Aphidoidea used in this book, compared with Heie (1980)

Heie (1980) This work

PHYLLOXEROIDEA APHIDOIDEA
Adelgidae Adelgidae
Phylloxeridae Phylloxeridae
APHIDOIDEA Aphididae
Pemphigidae Pemphiginae
Eriosomatinae Eriosomatini
Pemphiginae Pemphigini
Fordinae Fordini
Hormaphididae Hormaphidinae
Oregminae Cerataphidini
Hormaphidinae Hormaphidini
(Nipponaphidini) Nipponaphidini
Phloeomyzidae Phloeomyzinae
Thelaxidae Thelaxinae
Anoeciidae Anoeciinae
Mindaridae Mindarinae
Drepanosiphidae Drepanosiphinae
Drepanosiphinae Drepanosiphini
Phyllaphidinae Phyllaphidini
Chaitophorinae Chaitophorinae
Greenideidae Greenideinae
Greenideinae Greenideini
Cervaphidinae Cervaphidini
Aphididae Aphidinae
Pterocommatinae Pterocommatini
Aphidinae: Aphidini Aphidini
Aphidinae: Macrosiphini Macrosiphini
Lachnidae Lachninae
Lachninae Lachnini
Cinarinae Cinarini
Traminae Tramini

SYSTEMATIC TREATMENT OF APHID GENERA

(in alphabetical order)

ACANTHOCHERMES Kollar Phylloxeridae

Only one known species, with spiracles on abdominal segments 2-5 as in

Phylloxera and Viteus, but characterized by the curious star-like tubercles
borne by immatures. The adult apterae bear small spiculose tubercles on head,
thorax and abdomen. The anterior abdominal tergites each bear two pairs of
540 Acyrthosiphon

these tubercles, the spinal pairs being shorter than their basal width and the
lateral pairs being a little longer.

Acanthochermes quercus Kollar Recorded from Quercus robur and Q.

petraea in Sweden, Central Europe (Hungary, Germany) and Portugal. The
life cycle is remarkable in having only two generations per year, fundatrices
and sexuales. The fundatrices induce small ring-shaped galls on the upper
surface of the leaf, opening on the underside. Morphology and developmental
stages were illustrated by Grassi (1912) and Tavares (1931) gave an account
of the biology.

ACYRTHOSIPHON Mordvilko Aphidinae: Macrosiphini

A genus widespread in the palaearctic and northern oriental regions (85

species) and with another 15 species in North America. Most Acyrthosiphon
feed on herbaceous plants, particularly Leguminosae, Rosaceae and Euphor-
biaceae. None of the species in this genus host-alternate and associations with
trees are all secondary. In addition to the references in B & E, 1984, Richards
(1972b) and L.K. Ghosh (1986) give keys for Canada and Himachal Pradesh
respectively.

Acyrthosiphon aurlandicum Heikinheimo Apterae probably green in life,

2.1-2.5 mm, with siphunculi hardly longer than CAUDA, their apices slanted
outwards. Only recorded (apterae and oviparae) from Salix lapponum in
Norway (Heikinheimo, 1966). Alatae are unknown.

Acyrthosiphon elaeocarpi Tao Apterae whitish-yellow with yellowish-brown

siphunculi and blackish-brown antennae, apices of tibiae, tarsi, CAUDA and
anal plate; BL c. 3.5mm. On Elaeocarpus serratus in Taiwan (Tao, 1963).
Biology and sexual morphs unknown.

Acyrthosiphon evodiae (Takahashi) Apterae are yellow in life, about

2.3 mm, with black antennae and legs, brown-black siphunculi and a pale
CAUDA. Alatae have wings 'somewhat clouded along the veins'. Described
(in Macrosiphum) from specimens found feeding on young leaves of Euodia
triphylla in Taiwan (Takahashi, 1929). Life cycle unknown.

Acyrthosiphon gossypii Mordwilko (see B & E, 1984, p. 210). Green in

life, with adults as well as immatures dusted with fine wax (unlike A. pisum,
in which adult apterae have no wax bloom). Apterae 2.5-3.5 mm. On Cruci-
ferae, Leguminosae, Malvaceae and Zygophyllaceae in Central Asia, Middle
East, North Africa and India. Trees recorded from are Hibiscus esculentus,
Robinia pseudoacacia and Sophora japonica. Intraspecific variation in host
 Adeiges 541

plant preferences and life cycle are discussed by Müller and El Tigani (1986).
2n = 6.

Acyrthosiphon macrosiphum (Wilson) Yellow-green in life, with darker

antennae and tibiae, 1.5-1.9 mm. Holocyclic, with alate males, on leaves of
Amelanchier spp. in USA (California, Colorado, Utah, Oregon, Montana)
and Canada (British Columbia, Saskatchewan). Sambucus canadensis is also
listed as a host, but specimens have not been seen from that plant. References:
Palmer, 1952; Eastop, 1971. 2n = 10*.

Acyrthosiphon pisum (Harris) Pea Aphid (see B & E, 1984, p. 212) Apterae
rather large (2.5-4.4 mm), pale green or pink, usually in colonies on young
growth or pods of Leguminosae; only tree recorded from is Robinia pseu-
doacacia. 2n = 8.

Acyrthosiphon sophorae Narzikulov and Umarov Apterae rather large

(about 2.4 mm), pale green with waxy bloom, on Sophora spp. (alopecuroides,
japonica) in Kazakhstan and Tadzhikistan. Life cycle not known. References:
Narzikulov and Umarov, 1969; Eastop, 1971.

ADELGES Vallot Adelgidae

Distinguished by having five pairs of abdominal spiracles, whereas members

of the other genus of Adelgidae, Pineus, have only four distinct pairs. As in
Pineus the primary host of holocylic species is Picea, but the secondary hosts
are Abies, Larix, Pseudotsuga and Tsuga, never Pinus. The galls on Picea are
cone-like, often resembling miniature pineapples. The complete life cycle of
holocyclic species takes two years, and involves up to seven different morphs
(see p. 7). Several species are anholocyclic with a reduced number of morphs
and no host alternation, either living on Picea (in which case a gall may or
may not be formed by a 'pseudofundatrix'), or living on what was the original
secondary host.
The name Chermes was once commonly applied to Adelges, but is now
suppressed (Eastop, 1963). Some authors have used Adelges in a narrow sense
for the A. lands group and placed other species in Aphrastasia, Dreyfusia,
Cholodkovskya, Gilletteella and Sacchiphantes, but this generic classification
is based on rather slight differences and we use Adelges in its broader sense
to cover all these groups. Carter (1971) gave an account of the British species
and a check list of the world fauna, and Carter (1976) keyed the galls of British
Adelgidae on Sitka spruce. Other accounts are by Börner (1908), Börner
and Heinze (1957), Annand (1928: North America), Heinze (1962: Central
Europe), Binazzi and Covassi (1991: subgenus Dreyfusia in Italy), Inouye
(1953: Japan) and A.K. Ghosh (1984a: India).
542 Adeiges

Adeiges (Sacchiphantes) abietis (Linnaeus) Anholocyclic on Picea spp.

(abies, glauca, jezoensis, koyamai, sitchensis), with only two generations per
year; a pseudofundatrix that induces a pineapple-like gall (Fig. 123A), and
yellow to yellow-green alate gallicolae that leave the gall in late summer but
do not generally disperse far, often laying eggs on the same or neighbouring
spruce trees. The galls are hard, green and covered with trichomes; Rohfritsch
(1990) studied their development in detail. Ewert (1967) found differential
levels of infestation by A. abietis galls among spruce clones, Mitchell and
Maksymov (1977) studied predators, and Lasota et al. (1983) described a
heavy fungal attack on the galls. Found throughout Europe, North Africa
(Morocco), India and North America. Japanese records of abietis should pro-
bably be referred to A. japonicus, and most records from Larix are probably
of A. viridis. 2n = 18 (Steffan, 1968).

Fig. 123. Galls of Adelges on Picea. A, A. abietis on P. glauca (after Patch, 1910a); B, A. cooleyi on
P. sitchensis (after Carter, 1976); C, A. nordmannianae on P. orientalis; D, A. lands on P. sitchensis
(after Carter, 1976); E, A. pectinatae (after Cholodkovsky, 1907).
 Adeiges 543

Adeiges (Gilletteella) cooleyi (Gillette) Typically host-alternates between

Picea spp. (engelmanni, pungens, sitchensis) and Pseudotsuga spp. (macro-
carpa, menziesii). The galls on shoot tips of Picea are characteristically
elongate, with green, pink or red gall chambers and long needles protruding
from them (Fig. 123B). Gumming (1962b) described a monomorphic, non-
galling, anholocyclic form on P. glauca in Canada, and an anholocyclic
form also occurs on Pseudotsuga in both Europe and California, with alate
sexuparae developing in spring, but these are all gynoparae and produce only
sexual females, so that the sexual cycle is lost. Steffan (1970) applied the name
coweni Gillette to this latter aphid. Parry (1978a, b; 1980) and Parry and
Spiers (1982) have made extensive studies of the population biology of A.
cooleyi in Scotland, Teucher (1955) studied natural enemies, and Stephan
(1987) reported differences in resistance of Pseudotsuga provenances in Ger-
many. A. cooleyi occurs throughout Europe, North America and in Tasmania.
2n = 22 (Steffan, 1968).

Adelges (Aphrastasia) funitecta Dreyfus (= tsugae Annand) Host-alternating

between Picea polita and Tsuga sieboldii in Japan (Inouye, 1953); elsewhere
apparently anholocyclic on Tsuga spp. (canadensis, heterophylla, chinensis).
McClure (1989a) described sistens and apterous progrediens morphs and also
alate sexuparae from populations on Tsuga canadensis in Connecticut, USA;
the sexuparae migrated to Picea spp. and laid eggs, but sexuales failed to
develop, even on Picea polita, the recorded primary host in Japan. Recorded
from North America, India, Japan and Taiwan.

Adelges (Gilletteella) glandulae Zhang Host-alternating between Picea spp.

(brachytyla, likiangensis) and Abies ?faberi in Sichuan Province, China
(Zhang et al., 1980). The galls are ovate, cone-like, without associated needles.
Hiemosistens morph on Abies overwinters in second instar. This species is
described as anholocyclic, in spite of being gall-forming and host-alternating.
The fundatrix/?pseudofundatrix is undescribed.

Adelges (Dreyfusia) joshii Schneider-Orelli and Schneider Anholocyclic,

with three generations per year (hiemosistens and two aestivosistens) on Abies
pindrow in western Himalaya. Adults are dark brown, 0.7-1.2 mm, covered
with wax. Ghani and Rao (1966) give a detailed account of the morphology
and life cycle.

Adelges (Sacchiphantes) karafutonis Kono and Inouye Galls of this species

on young shoots of Picea jezoensis are small, with only 5-15 scales (Inouye,
1953). Only the gallicolae are described and the secondary host (presumably
a Larix species) is unknown. For differences from the closely related A.
torii see Eichhorn and Carter (1978). Recorded from Sakhalin and Japan
(Hokkaido).

Adelges (Dreyfusia) knucheli Schneider-Orelli and Schneider Host-alter-

nating between Picea smithiana and Abies spp. (pindrow, spectabilis) in
544 Adeiges

western Himalaya (Pakistan and Kashmir), or anholocyclic on Abies pindrow

where the primary host is absent, e.g. in Murree Hills of Pakistan. The pine-
apple- or cone-like galls on P. smithiana vary considerably in size and shape.
The sistens generations on Abies pindrow are dark brown, with wax filaments
radiating in a star-like fashion, and occur on main stems of older firs as well
as on branches of younger trees. Progrediens apterae and alate sexuparae
develop on new-grown needles, the latter without secreting wax. Ghani and
Rao (1966) provided detailed descriptions of all stages and morphs. Predators
were discussed by Pschorn-Walcher (1964). [Stebbing's (1910) account of
Chermes himalayensis probably applies to this adelgid; but see Schneider-
Orelli and Schneider (1954).]

Adeiges laricis Vallot (= strobilobius Kaltenbach), Plate la, b The oldest-

named member of a taxonomically very difficult complex of adelgids which
have host alternation between Picea and Larix, or have an incomplete cycle
restricted to either the primary or the secondary host. Galls of laricis sensu
stricto on Picea are typical of the group (Fig. 123D); they are globular or
ovate, with a cream or ivory coloration and waxy texture, becoming pink or
brown just prior to opening. They may be terminal, or have some growth
of the shoot beyond the gall. Gall development was described in detail by
Rohfritsch (1990). Predation within galls by dipterous larvae was described by
Mitchell and Maksymov (1977). Gallicolae of A. laricis migrate to Larix in
June-July (Fig. 1); they are 1-2 mm, blackish, and secrete little or no wax. The
hiemosistens generation on Larix in spring is dark grey without wax, but the
progrediens apterae left after departure of the sexuparae in June-September
produce abundant wax and honeydew; these deposits from a heavy infestation
cause the foliage to turn blue (Börner and Heinze, 1957; Carter, 1971).
Adelges laricis sensu stricto occurs in Europe and North America and is
recorded from Picea abies, koraiensis, mariana, rubens, sitchensis (primary
hosts), and from Larix decidua, x eurolepis, laricina, leptolepis (secondary
hosts). 2n = 20 (Steffan, 1968).
Other members of the laricis group, some of which were only described
from one or two morphs and are possibly not distinct species, are as follows:
aenigmaticus Annand Only the apterous ? progrediens morph is described
by Annand (1928). It was collected on Larix laricina in Maine, USA, and
distinguished from laricis by its smaller, rounded dorsal sclerites bearing
round, separate wax pores, and by the short and thick third antennal seg-
ment. Similar specimens were collected by F.W. Quednau on the same
host in Quebec, Canada (F.W. Quednau, pers. comm., 1971). Life cycle is
unknown.
diversis Annand Progrediens apterae collected on cones of European larch
(Larix decidua) in Oregon, USA, were thinly covered with short wax threads
(Annand, 1928). Only the first instar ?sistens, progrediens and immature
?sexupara are described. The progrediens is distinguished by having no
sclerites on abdominal tergites 5-8 and only small sclerites with few wax pores
on the more anterior tergites. Not recorded since the original description.
 Adeiges 545

isedakii Eichhorn Host-alternating between Picea jezoensis (var. hondoensis)

and Larix leptolepis in Japan (Eichhorn and Carter, 1978). The gall on Picea
is similar to that of laricis, but larger. The gallicola has a red body and
greenish wings, is smaller (1.0-1.5 mm) than the non-migratory gallicola of
A. japonicus (see below), and has no wax glands on the head. Only the first
instar hiemosistens (neosistens) has been described from Larix.
japonicus Monzen Anholocyclic, pseudofundatrices inducing cone-like galls
on shoot tips of Picea jezoensis and P. sitchensis in Japan and Sakhalin
(Inouye, 1953; Pashchenko, 1988b). Gallicolae emerging from galls in August-
September are large (1.9-2.8 mm) and with or without cephalic wax glands
(Eichhorn and Carter, 1978); some do not disperse, so that individual trees
may develop heavy levels of infestation (Kamata, 1986).
karamatsui Inouye Anholocyclic on secondary hosts (Larix leptolepis,
gmelinii, koraiensis) in Japan, Sakhalin and Korea. Sistens, apterous pro-
grediens and sexupara are described (Inouye, 1953). Sexuparae, small and
dark brown to black with abundant wax, fly to Picea in Japan, but die or
produce eggs which do not hatch (Eichhorn and Carter, 1978); an analogous
situation to that with A. funitecta on Tsuga in USA.
lapponicus (Cholodkosky) Anholocyclic, pseudofundatrices inducing galls
on both native and exotic Picea spp. Dark red gallicolae about 1.6mm long
emerge from galls in June-July and lay reddish eggs on needles of Picea
with little or no secretion of wax. In northern and Central Europe, and
more recently recorded from Kirghizia (Gabrid, 1981). Differential infes-
tation of Picea spp. in northern Finland was studied by Häggman and Rousi
(1986).
lariciatus Patch Host-alternating between Picea spp. (abies, glauca, mariana,
pungens) and Larix spp. (laricina, lyalli) in North America (Alberta, Manitoba,
Saskatchewan, Utah). Galls are globular, often subterminal and to one side
of shoot, green when young to red and brown when mature. Cumming (1968)
gave a detailed account of morphs and life cycle in Canada.
oregonensis Annand Only known from the original description of sistens and
progrediens morphs from Larix occidentalis in northwestern USA (Oregon,
Washington, Montana). Differences from laricis were discussed by Annand
(1928). It feeds on the needles and on the twigs at the needle bases. The
primary host, if any, is unknown.
pontaninilaricis Zhang Host-alternating between Picea retroflexa and Larix
pontaninii in Sichuan Province, China. Described as a subspecies of laricis on
the basis of differences in number and distribution of wax glands (Zhang
et al., 1980).
tardoides (Cholodkovsky) Host-alternating between Picea abies and Larix
sibirica in northeastern Europe. The gallicolae migrate to the secondary host
in July-August, later than laricis, and unlike those of laricis, secrete abundant
wax (Börner and Heinze, 1957, p. 346).
546 Adeiges

tardus (Dreyfus) Anholocyclic on Picea spp. (abies, glauca, obovata, pun-

gens) throughout continental Europe. The gallicolae emerge from late June
to September, later than lapponicus, and lay eggs on exposed parts of the trees
under a covering of white wax. Due to non-dispersal of the gallicolae, the galls
may become very numerous and can severely affect growth of young spruce
trees. Baurant (1968) studied the phenology of A. tardus in relation to Picea
abies in Belgium, and suggested that its synchronization with the host might
be upset by selection of appropriate spruce varieties. 2n = 20 (Steffan, 1968).

Adelges (Dreyfusia) merkeri Eichhorn Host-alternating between Picea orien-

talis and Abies alba, with a rather restricted distribution in Europe (Germany,
southern Sweden, one record from Austria); origins unknown. Galls on Picea
resemble those of A. nordmannianae but are usually larger (7-22 mm) and
open in June-July (Eichhorn, 1975). On Abies, colonies are found on the
trunk and crown region of older firs, or on all parts of young and seedling
trees, where damage may be severe with swelling and stunting of infested
branches, especially at their bases. As well as the hiemosistens there are one
or even two aestivosistens generations. Adult sistentes generally have less
wax wool than either nordmannianae or piceae. Pschorn-Walcher and Zwölfer
(1960) provided ecological notes and Eichhorn (1968) studied population
dynamics on Abies, especially the influence of natural enemies.

Adelges (Dreyfusia) nordmannianae (Eckstein) (= nüsslini C.B.) Host-

alternating between Picea orientalis and Abies spp. (alba, bornmuelleriana,
cilicica, nordmanniana, etc.). Believed to be endemic and relatively uninju-
rious in the Caucasus and East Pontus mountains (former USSR and Turkey),
where there are mixed stands of Picea orientalis and Abies nordmanniana,
but causing severe damage to Abies where non-endemic in Europe, North
America, New Zealand (see Zondag, 1982) and Tasmania. Galls on Picea
(Fig. 123C) are 2-15 mm long, terminal, globular, pinkish and resembling
strawberries when young, later becoming greenish with red or purple colora-
tion at bases and tips of scales. They mostly open in June (Eichhorn, 1975).
There is never more than one aestivosistens generation and even this is
lacking in some populations. On Abies alba there appear to be two forms of
A. nordmannianae: 'typica' which infests branches, twigs and needles of young
trees in sunny dry locations - the attack of the progrediens generation causing
severe deformation of needles and shoots and often killing the tree (see Varty,
1956); and form 'schneideri' which infests the trunks of older firs 30-120 years
old in forests (studied by Pschorn-Walcher & Zwölfer, 1958; and Eichhorn
and Pschorn-Walcher, 1972). The latter form would not transfer to young
trees (Schneider-Orelli et al., 1929). Colonies of sistentes on stems are black
with a grey wax bloom and only a fringe of wax wool (cf. A. piceae). Eichhorn
(1969a) studied the natural enemies of A. nordmannianae in Turkey, and also
worked on morph determination (Eichhorn, 1969b), life cycle (Eichhorn,
1991) and embryonic development (Eichhorn, 1970). Most recent ecological
studies of this species have been in Austria (Sturzer-Gilbert, 1982; Bauer-
Schmid, 1983). 2n = 22 (Steffan, 1968).
 Adeiges 547

Adeiges (Aphrastasia) pectinatae (Cholodkovsky) Host-alternating between

Picea (abies, obovata) and Abies spp. (alba, balsamea, concolor, etc.), in
northern Europe (Sweden, Finland, Latvia, Ukraine) and east to Siberia.
Recently recorded from eastern Norway on ten species of Abies, although the
gall generations on Picea were not found there (Austará, 1990). The galls
(Fig. 123E) are compact, cone-like, dull green turning reddish-brown before
opening, about 10-17 mm long. Reddish-brown gallicolae fly to Abies in June-
July. Feeding on Abies is restricted to the needles. Frolowa (1924) studied
cytology of egg and sperm maturation in this species. 2n = 20 (Steffan, 1968).
Populations alternating between Picea spp. (glehni, jezoensis, etc.) and
Abies (sachalinensis, veitchii) in Japan are regarded as a separate subspecies,
A. pectinatae ishiharai Inouye (Eichhorn and Carter, 1978). Its morphs and
life cycle are described in detail by Inouye (1953). Fang (1981) described the
pest status of what is possibly this subspecies on Picea jezoensis, P. koraiensis
and Abies nephrolepis in China.
An anholocyclic form in the pectinatae group, Adelges pindrowi Yaseen
and Ghani, occurs on Abies pindrow in Pakistan. Information about its life
cycle is provided by Yaseen and Ghani (1971).

Adelges (Dreyfusia) piceae (Ratzeburg) Balsam Woolly Aphid Anholo-

cyclic on Abies spp. (alba, balsamea, cilicica, fraseri, etc.), with one hiemo-
sistens and usually one aestivosistens generation. In favourable conditions
a second, or even a third, aestivosistens generation may occur (Pschorn-
Walcher, 1964). In North America there is sometimes also a generation of
progrediens apterae and alatae. On A. alba in Europe this adelgid normally
infests the trunk and larger branches of older trees (30-100 years) and is
of little economic importance. Introduced populations in North America,
however, attack stems and buds of the crown region of Abies balsamea and
A. fraseri, causing extensive injury. Young trees of A. balsamea are killed by
'gout disease' (excessive shoot swellings near the buds); heavy stem attack may
give rise to severe transpiration stresses by the formation of rotholz or com-
pression wood (Balch, 1952), causing crown dieback and a degrade of the
timber (Carter, 1971). Hain et al. (1983) studied the host's defence reaction
against stem attack.
The economic importance of A. piceae has resulted in an extensive
literature; Schooley and Oldford (1981) provided an annotated bibliography,
and Johnson and Lyon (1988) gave a short, well-illustrated account. Biological
control measures up until 1958 were summarized by McGugan and Coppel
(1962). Foottit and Mackauer (1983) used morphometric studies to distinguish
and key three forms of A. piceae in North America, thought to represent
separate introductions. They designated these as subspecies (p. piceae, p.
canadensis and p. occidentalis), although it is questionable whether the
subspecies category is appropriate for anholocyclic species. The form known
as canadensis is similar or identical to populations of piceae causing compres-
sion wood in Abies grandis in Scotland (Busby, 1964). Binazzi and Covassi
(1991) differentiated an anholocyclic population on Abies nebrodensis in
Italy as a new species, A. (D.) nebrodensis. 2n is unknown.
548 Adeiges

Adeiges (Dreyfusia) prelli (Grosmann) Host-alternating between Picea

orientalis and Abies nordmanniana or (rarely) Abies cephalonica. Like A.
nordmannianae, A. prelli is indigenous to Turkey (Eichhorn, 1969a) and
introduced into Central and Western Europe. The galls are bluish-green, larger
than those of A. nordmannianae (6-30mm long) and open later, in July-
August (Eichhorn, 1975). On Abies nordmanniana in Western Europe a
hiemosistens and one or two aestivosistens generations occur, almost exclu-
sively attacking the crown region of mature trees (Eichhorn, 1956). The
hiemosistens on young shoots is more sclerotized and has less wax than that
on stems or branches (Eichhorn, 1964). North American populations of
A. piceae on Abies balsamea closely resemble A. prelli, which has not however
been positively identified from that host (or from Abies alba).

Adelges (Dreyfusia) todomatsui (Inouye) Anholocyclic on Abies sachalinen-

sis (incl. var. mayriana) and A. veitchii in Hokkaido, Japan, with a hiemo-
sistens and one or possibly two aestivosistens generations (Eichhorn and
Carter, 1978). No progrediens (needle-dwelling) morph has been found.
Inouye (1953) observed that two biological forms occur that are indistin-
guishable morphologically: a stem-dwelling form on young and mature (pole-
stage) trees, and a bud-dwelling form that lives under the scales of the previous
year's shoots and causes gall-like enlargement of buds. Predators were studied
by Pschorn-Walcher (1964).

Adelges (Sacciphantes) torii (Eichhorn) Host-alternating between Picea

jezoensis var. hondoensis and Larix leptolepis in Honshu, Japan. Galls are
small and reddish, similar to those of A. karafutonis, opening in August.
Eichhorn and Carter (1978) provided a detailed account.

Adelges (Cholodkovskaya) viridana (Cholodkovsky) Anholocyclic on Larix

spp. (decidua, x eurolepis, gmelinii, leptolepis, sibirica) in Europe (Sweden,
UK, France, Germany, Italy, Ukraine) and east to China, Korea and Japan,
although somewhat sporadic or local in occurrence. The life cycle is variable,
probably depending on latitude and/or temperature. The overwintering first
instar sistens is yellowish-green to green and usually under bark on the stem.
The large (1.8-2.5 mm) adult sistentes, yellowish-green and surrounded by wax
secretion, occur in May-June. Their eggs develop either as overwintering
hiemosistentes or as alate progredientes; first instars of the latter morph are
very active and move to tender young extension shoots to feed at the needle
bases. The adult alate progrediens is quite large (2.0-2.6 mm), grey-green and
mostly non-migratory, laying eggs on the needles. These mostly develop as
hiemosistentes but, in France and Germany (Gaumont, 1954; Steffan, 1964),
some of them develop into a second generation of alate progredientes, and
in suitable conditions these may even give rise to a third partial alate pro-
grediens generation. 2n = 24 (Steffan, 1968).
In northern Russia, populations of the closely-related A. (Ch.) viridula
(Cholodkovsky) occur on Larix sibirica, and these seem always to have a single
annual (hiemosistens) generation, no alatae being known. Its overwintering
 Aiceona 549

first instars have fewer wax pores on posterior abdominal segments than
A. viridana.

Adelges (Sacchiphantes) viridis (Ratzeburg) Host-alternating between Picea

spp. (abies, koraiensis, orientalis, sitchensis, etc. - see Weis, 1955), and
Larix spp. (decidua, x eurolepis, gmelinii, leptolepis). Steffan (196la) made
a detailed study of this adelgid in Germany (but see also Eichhorn, 1989). The
pineapple galls on Picea are similar to those of A. abietis but open earlier,
in July or early August. Miszta (1987) has studied their volatile oil composition
in comparison to normal plant tissue. Immatures within the gall, and the alate
gallicolae when they emerge, are reddish-yellow to brown (cf. A. abietis).
Hiemosistens immatures and adults are green or brown. Sexuparae developing
on Larix the following spring are pale greenish with sparse wax and their
feeding causes yellowing and kinking of the newly-grown needles (Carter,
1971). The return migration to Picea is quite early, in May-June. Adelges
viridis occurs throughout Europe and is also recorded from China, where it
migrates from Picea koraiensis to Larix gmelinii (Fang et al., 1983). 2n = 18
(Steffan, 1968).
Apart from A. abietis, anholocyclic on Picea, there are two other described
taxa that are closely related to A. viridis, but only known from Larix:
kitamiensis Inouye Attacking Larix leptolepis in Hokkaido, Japan, some-
times so heavily that the entire trunk of a 10-20-year-old tree is covered in
its white wax wool. Hiemosistens immatures and adults are dirty blue in colour
(cf. viridis). The small (about 0.6-0.7 mm) sexuparae maturing in May-June
leave Larix for an unknown primary host; no galls have been found on Picea
abies, even where this tree occurs in mixed stands with L. leptolepis. For
further details and differences from viridis see Inouye (1963).
segregis Steffan Anholocyclic on Larix decidua in Central Europe, with two
generations per year, one alate and the other a 'pseudohiemosistens'. The
overwintering first instar larva of the latter has a shorter rostrum than the
neosistens of viridis (Steffan, 196la).

AICEONA Takahashi Anoeciinae

Aiceona is not as closely related to Anoecia as might be thought from its name
and placement in Anoeciinae. There are about 14 recognized species, all in
East and Southeast Asia, mostly associated with Lauraceae. Several species
are described from unidentified hosts. Unusually, oviparae as well as males
are alate, and they may be found in colonies at almost any time of year.
Siphunculi are well developed as hairy cones in all female morphs, but
strangely absent in males. Accounts of Aiceona are available for India (M.R.
Ghosh and Raychaudhuri, 1973), northeast India (Raychaudhuri et al.,
1980d) and Japan (Takahashi, 1960a). The genus is clearly in need of revision.
550 Aiceona

Aiceona actinodaphnis Takahashi Apterae are 2-3 mm, purplish-black

dusted with white wax powder, living under leaves and on new growth of
various Lauraceae (Actinodaphne, Cinnamomum, Lindera, Litsea). Alatae
have dark-veined smoky wings with a pale area between the tip of the
pterostigma and Rs. Sexual morphs are unknown. Recorded from China,
Okinawa and Japan (Tseng and Tao, 1938). A related species with paler
wings, A. siamensis, is described from unidentified Lauraceae in Thailand
(Takahashi, 1941).

Aiceona himalaica Miyazaki Appearance in life unrecorded, apterae 2.4-

2.8 mm; cleared and mounted apterae have a pale body contrasting with the
dark hind tibiae. Apterous viviparae, alate oviparae and males are described.
Collected in July in Nepal, from Engelhardtia spicata (Juglandaceae), an
unusual host association for this genus (Miyazaki, 1977).

Aiceona japonica Takahashi Apterae are about 3 mm, blackish-brown,

covered with white wax powder, on leaves and young growth of Lauraceae
(Actinodaphne lancifolia, Cinnamomum camphora, ?Litsea sp.) in Japan,
and also recorded from Korea (Paik, 1972). The wings of the alata are
uniformly dusky without any clear area. Alate oviparae and males occur
along with apterous and alate viviparae in May (Takahashi, 1960a), although
Miyazaki (1977) expressed some doubt about Takahashi's description of the
ovipara of this species.

Aiceona malayana Takahashi Apterae pale greenish-yellow, 2.2-2.5 mm,

attacking young leaves of ?Actinodaphne sp. near Kuala Lumpur, Malaya.
Alatae have wings clear or only faintly dusky. Distinguished by its short
antennal hairs, this species is only known from the original collection, when
it was recorded erroneously as A. osugii (Takahashi, 1950, 1960a). Sexual
morphs and life cycle are unknown.

Aiceona parvicornis Miyazaki Appearance in life is unknown; mounted

apterae (about 2.7 mm) have dark head, prothorax, coxae and femoro-tibial
joints. Alatae have dusky wings. Collected in May from Lindera pulcherrima
in Nepal (Miyazaki, 1977). Sexuales and life cycle are unknown.

Aiceona pseudosugii David, Sekhon and Bindra Apterae are 2-3 mm, pale
yellow with dark hind tibiae, occurring in small colonies on undersides of
young leaves, with attendant ants. Alatae have unpigmented wings. The
only identified host is Machilus odoratissima (David et al., 1970). Aiceona
longisetosa M.R. Ghosh and Raychaudhuri, 1973 seems to be a synonym.
Recorded from Himachal Pradesh and West Bengal, India. Sexuales and
life cycle are unknown.

Aiceona retipennis David, Narayanan and Rajasingh Apterae are 2.5-

2.8mm, yellowish-white, on undersides of leaves and young shoots (L.K.
Ghosh, 1972). Alatae have very dark wings with a clear patch between the
 Aleurodaphis 551

pterostigma and Rs like that in A. actinodaphnis. Other distinguishing fea-

tures of this species are the very long fine antennal hairs and short last rostral
segment (R IV+V) only 0.6-0.7 times HT II. Pal and Raychaudhuri (1977)
describe the alate male, collected in February. Its only identified hosts in the
Lauraceae are Machilus gamblei (BMNH colln, 1 aptera leg. K. Narayanan)
and Machilus sp. (Chakrabarti et al., 1988); the various records from plants
in other families seem doubtful. Widely distributed in northern India. 2n= 18
(Khuda-Bukhsh, 1980).

Aiceona robustiseta M.R. Ghosh and Raychaudhuri Apterae are 2.0-

2.5 mm, creamy-yellow with a light dusting of wax, in colonies on young
growth attended by ants (M.R. Ghosh and Raychaudhuri, 1973). Alatae
have unpigmented wings. Alate oviparae, collected in February and June,
are described by Pal and Raychaudhuri (1977). Only known from Litsea
polyantha in West Bengal, India.

Aiceona titabarensis (Raychaudhuri and A.K. Ghosh) (= litseae A.N. Basu

and Hille Ris Lambers), Plate 5c, d Apterae are 2.1-2.7 mm, dirty brownish
to grey, dusted with wax powder, often heavily infesting the undersides
of leaves and young shoots of Litsea polyantha (A.N. Basu and Hille Ris
Lambers, 1968), with or without attendant ants. Alatae have smoky wings
as in A. actinodaphnis and A. retipennis, but the clear area adjacent to the
pterostigma is narrower and less clearly defined. The species was originally
described (in Lachnus) from Heteropanax fragrans (Araliaceae), a host which
is unusual enough to need confirmation. Alate oviparae can be found in
colonies with apterous and alate viviparae at least from June to December
(M.R. Ghosh and Raychaudhuri, 1973). Recorded from Sikkim and West
Bengal, India, and from Hong Kong and Vietnam (BMNH colln).

ALEURODAPHIS van der Goot Hormaphidinae: Cerataphidini

A little-known genus with four or five nominal species in eastern Asia. It has
aleyrodiform apterae with a complete crenulate margin of wax glands, as in
Cerataphis, but distinguished from that genus by absence of frontal horns and
a more evident division between prothorax and mesothorax. Life cycles are
unknown, but an undescribed species has been collected (by G. Zhang and
VFE) from boat-shaped galls on Sinojackia xylocarpa at Hangzhou, China.
Others are described from Compositae (three species), Bambusa and Salix.
Two of the species on Compositae in Japan are monoecious and anholocyclic
(Takahashi and Sorin, 1958).

Aleurodaphis antennata Chakrabarti and Maity Only the aleyrodiform

aptera (BL 1.3-1.5 mm) is described, from Bambusa sp. in Uttar Pradesh,
552 Allocotaphis

India (Chakrabarti and Maity, 1982). The last rostral segment (R IV+V) is
shorter than in other Aleurodaphis (1.1-1.5 x HT II). Appearance in life is
unknown, presumably dark with marginal wax.

Aleurodaphis sinisalicis Zhang Only the aleyrodiform aptera (BL about

1.4 mm) is described, from Salix sp. in China (Zhang and Zhong, 1982b). The
host plant is unusual enough to require confirmation. Very similar, if not
synonymous with, A. blumeae van der Goot. Appearance in life and biology
unknown.

ALLOCOTAPHIS Börner Aphidinae: Macrosiphini

A genus for the one species, A. quaestionis (Börner), a rather large green aphid
rolling apple leaves in spring and migrating for the summer to Senecio doro-
nicum. In Central Europe, with a boreo-alpine distribution (Shaposhnikov,
1951; Hille Ris Lambers and Wildbolz, 1958; B & E, 1984, p. 215).

ALLOTHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini

The apterae of the only known species have long marginal prosomal hairs like
Thoracaphis, but lack siphuncular pores and marginal hairs on the reduced
abdominal tergites 2-7. They have a less sclerotic dorsum than Thoracaphis,
with an irregular mosaic-like ornamentation.

Allothoracaphis piyananensis Takahashi Apterae are pale yellow to yellow-

ish-brown, flattened dorsoventrally, on the undersides of leaves along the
main veins. Recorded from Quercus glauca and Q. morii in Japan and Taiwan
(Takahashi, 1958a). The alate morph is unknown, and the species is appar-
ently anholocyclic.

ALLOTRICHOSIPHUM Takahashi Greenideinae: Greenideini

A genus for two oak-feeding species, narrow-bodied and with long siphunculi
like Eutrichosiphum, but differing in the presence of distinctly capitate or
spatulate hairs.
 Anoecia 553

Allotrichosiphum assamense. Rauchaudhuri, Ghosh, Banerjee and Ghosh

Apterae about 1.6mm, appearance in life unrecorded but probably green
or pale brown; known only from one apterous vivipara and one immature
collected on Quercus dealbata in Assam, India (Raychaudhuri et al., 1973).
(Note that this is a different species from Eutrichosiphum assamense.)

Allotrichosiphum kashicola (Kurisaki) Apterae (fundatrices) 2.3-2.8 mm,

colour in life unrecorded, probably green or pale brawn. Alatae dark greenish-
brown, clear-winged, with siphunculi nearly as long as body. On young foliage
of Quercus spp. (acuta, glauca, myrsinaefolia) in Honshu and Kyushu, Japan.
The life cycle is greatly abbreviated, with sexual morphs (greenish alate
oviparae and yellow alate males) appearing in May. The rather flat eggs are
laid on the undersides of leaves. At Osaka there are only two generations per
year (fundatrices and sexuales), although at Tokyo additional parthenogenetic
generations have been observed (Takahashi and Sorin, 1957, 1959). Apterous
viviparae other than fundatrices have not been detected.

ALOEPHAGUS Essig Pemphiginae: Fordini

A genus with only one known species, probably of African origin (Hille Ris
Lambers, 1954a).

Aloephagus myersi Essig Emigrant alatae suspected to be this species, BL

about 1.8mm, have been collected from galls on Pistacia sp, (presumably P.
ethiopica) in Kenya (D. Hille Ris Lambers, pers. comm.; specimens in BMNH
colln). Presumed to host-alternate in Africa between Pistacia and Aloe, where
it forms densely wax-dusted, ant-attended colonies under the leaf-bases.
Anholocyclic on Aloe in Europe and North America (B & E, 1984, p. 215).
The galls and the emigrant alatae are as yet undescribed. 2n = 22.

ANOECIA Koch Anoeciinae

A distinctive holarctic genus of uncertain taxonomie position, with about 20

species, many of which are poorly known. Some have host alternation from
Cornus to roots of Gramineae (in one case, Onagraceae). Others are entirely
subterranean on Gramineae or Cyperaceae. The alate sexuparae, distinguished
by their dark posteriodorsal abdominal patch and the large black pterostigmal
spot on the forewing, are a common sight on Cornus leaves in the autumn,
alighting and depositing ant-attended clusters of small, yellow and/or brown
sexual morphs. The oviparae lay their eggs on the bark of the trunk. Different
554 Anoecia

species utilize the native Cornus in North America, Europe and eastern Asia.
This is a difficult group in need of revision; in the BMNH collection there
are undescribed species from Cornus mas in Italy, and from Cornus spp. in
Korea. See also B & E, 1984, pp. 217-218.

Anoecia corni (Fabricius), Plate 5a, b Host-alternating between Cornus

sanguinea and roots of Gramineae in Europe; anholocyclic on grasses and
some cereals in eastern Asia and North America. Apterae in spring on Cornus
are dark brown; fundatrices have reduced, 3-faceted eyes and 5-segmented
antennae, whereas subsequent generations have large compound eyes and
6-segmented antennae. Both spring migrant alatae and the sexuparae returning
to Cornus in autumn have a dark patch covering abdominal tergites 3-7. BL
of alata 1.9-3.0mm. See Knechtel and Manolache (1943) and Heie (1980)
for further information. 2n = 6.

Anoecia cornicola (Walsh) Host-alternating between Cornus spp. (amo-

mum, stolonifera) and roots of Gramineae in North America, and also
recorded from Brazil (BMNH colln) where it is presumably anholocyclic on
grass roots. Alate sexuparae have a dark dorsal abdominal patch but this is
apparently not present in spring migrants (Palmer, 1952). BL of alata 2.0-
2.5 mm. 2n= 10.

Anoecia fulviabdominalis (Sasaki) Host-alternating between Cornus spp.

(brachypoda, controversa) and roots of Gramineae in Japan; also anholocyclic
on roots of Gramineae and a major pest of upland rice in East and Southeast
Asia. Both spring migrant alatae and the returning sexuparae have dark dorsal
abdominal patches. Tanaka (1961) studied its biology in Japan. BL of alata
1.8-2.5 mm.

Anoecia major Börner Not yet found on Cornus in nature, but Börner
(1950) transferred sexuparae to C. sanguinea and obtained two-three genera-
tions of fundatrigeniae. Occurs widely in Europe on roots of Phalaris arun-
dinacea, and also recorded from Brachypodium pinnatum. BL 2.3-3.2 mm.
See also Heie (1980). 2n = 8.

Anoecia oenotherae Wilson Host-alternating between Cornus spp. (amo-

mum, paniculata, stolonifera) and roots of Oenothera biennis (Onagraceae)
in North America. Alatae are small (1.4-1.7 mm) and never have a solid patch
on abdominal tergites 3-7, but sexuparae may have dark cross-bands on these
tergites. See also Palmer (1952).

Anoecia vagans Koch Host-alternating between Cornus sanguinea and roots

of Gramineae in Europe. All the progeny of fundatrices on Cornus are spring
migrant alatae which, unlike those of A. corni, have no dark posterior
abdominal patch. The sexuparae returning to Cornus in autumn do, however,
have such a patch. BL of alata 2.3-3.9mm (Heie, 1980). 2n=12.
 Anomalosiphum 555

ANOMALAPHIS Baker Greenideinae: Cervaphidini

A genus for two Australian species living on Myrtaceae (one on Agonis and
the other on Leptospermum). The antennae of both apterae and alatae are
5-segmented, and abdominal tergites 7 and 8 each bear slender hair-bearing
processes. The siphunculi are cylindrical or tapering, with a ring of 3-8 hairs
on the distal half, and the CAUDA is broadly rounded.

Anomalaphis casimiri Carver Apterae are small (BL 1.3-1.7 mm) and
brown. Distinguished from the only other described species (A. comperei
Pergande) by the presence of polygonal reticulation on the siphunculi distal
to the ring of hairs, and by dorsal hairs of aptera mostly arising from
tubercles. Apterae and alatae occur together with intermediate forms on
Leptospermum sp. var. Red Damask in New South Wales, Australia (Carver,
1971). Appearance of colonies in life and biology are not recorded. (Ano-
malaphis comperei on Agonis has apterous oviparae and is presumably
monoecious and holocyclic.)

ANOMALOSIPHUM Takahashi Greenideinae: Cervaphidini

Six species associated with woody Connaraceae, Leguminosae and Poly-

galaceae in East and Southeast Asia. The hosts are often woody shrubs or
climbers, and the host of one species in China (A. takahashii Tao) is unknown.
The apterae have 4-segmented antennae, bear slender hair-bearing processes
on abdominal tergites 7 and 8, and have a CAUDA tipped with a stylus. Alatae
have 5-segmented antennae, much smaller processes on the posterior tergites,
and the caudal process often indistinct. Martin and Agarwala (1994) revised
the genus, describing three new species, two of them from shrubby species
of Dalbergia.

Anomalosiphum indigoferae Ghosh, Ghosh and Raychaudhuri Apterae in

life dark brown to black, BL 1.1-1.4 mm, described from young twigs of
Indigofera sp. in India (A.K. Ghosh et al., 1971e). Alatae are larger, BL
1.4-1.7 mm. Recorded fom Sikkim and West Bengal. Life cycle unknown.

Anomalosiphum murphyi Martin and Agarwala, Plate 12a, b Colour in life

unrecorded, apterae probably dark-pigmented; BL 1.2-1.6 mm. On Rourea
sp. and Xanthophyllum stipitatum in Singapore, and on ?Sindora sp. in
Sarawak. Alatae in Singapore in January. Life cycle unknown (Martin and
Agarwala, 1994).
556 Anuraphis

Anomalosiphum pithecolobii Takahashi Apterae probably green in life:

BL 1.3-1.4 mm. Alatae greenish-yellow. On young leaves and shoots of
Pithecellobium lucidum in Taiwan. Trapped alatae have also been seen from
Papua New Guinea. Life cycle unknown.

Anomalosiphum tiomanensis Martin Apterae pale green; BL 1.3-1.5 mm.

On new apical growth of Dalbergia torta in western Malaya. Apterae and
alatae collected in February. Life cycle unknown.

ANURAPHIS Del Guercio Aphidinae: Macrosiphini

A small palaearctic genus of plump-bodied, dull olive-green to brown aphids

with rather short appendages, typically producing 'pseudogalls' by crumpling
or rolling the leaves of Pyrus spp. in spring. The progeny of the fundatrix are
all alate and migrate to subterranean parts of Umbelliferae or Compositae.
For further information see B & E, 1984, p. 219. Heie (1992) gave an account
of the two species in northwest Europe.

APHANOSTIGMA Börner Phylloxeridae

A genus differing from Phylloxera in the absence of spiracles from abdominal

segments 2-5. Two species are described from Pyrus; A. piri (Cholodkovsky)
recorded from southern Europe, Ukraine, Georgia, Crimea, Israel, Lebanon
and Thailand (BMNH colln); and A. iaksuiense (Kishida), only known from
Japan. See B & E, 1984, p. 220.

APHIDOUNGUIS Takahashi Pemphiginae: Eriosomatini

A monotypic genus in Japan characterized by the single developed tarsal claw

of apterae on the secondary host (B & E, 1984, p. 373, as Watabura nishiyae).
Emigrant alatae from galls on Ulmus have short antennae, forewings with
unbranched media and small siphuncular pores without associated hairs.

Aphidounguis mali Takahashi Host-alternates between Ulmus parvifolia

and roots of Malus and Cydonia in Japan. Alatae have also been collected from
U. parvifolia in China (Hangzhou), and from U. japonica and U. mandshurica
 Aphis 557

in Korea. The Ulmus galls consist of several rolled leaves on one shoot;
initially only one of these is occupied by the fundatrix, but later its progeny
migrate to fill the empty leaf-rolls (Akimoto, 1983). Alate emigrants, BL
1.5-1.7 mm, have dark brown head, thorax, legs and antennae, and dark
bands between the abdominal tergites. Apterae on roots of secondary hosts
are small white aphids secreting wax wool.

APHIDURA Hille Ris Lambers Aphidinae: Macrosiphini

A palaearctic genus of about 12 species in the Mediterranean region, Eastern

Europe and Central Asia. Most of them are recorded from Caryophyllaceae,
but one species occurs in Central Asia on Prunus spp. of the microcerasus
group. Aphidura are Myzus-like aphids but have a pair of mesosternal mam-
mariform processes.

Aphidura bozhkoae (Narzikulov) Shiny black aphids, BL about 2mm,

infesting apical leaves of Prunus spp. (erythrocarpa, incana, ulmifolii, ver-
rucosa, but not avium or cerasus) in Central Asia (Tadzhikistan, Kazakhstan,
Kirgizia, Georgia) (Narzikulov, 1958, 1965b). Life cycle is unknown.

APHIS Linnaeus Aphidinae: Aphidini

Of the more than 400 species of Aphis, only about 50 are recorded from trees.
Some of these are monoecious on particular tree species, some host-alternate
between trees and herbs, and some are polyphagous. For an introduction to
the genus see B & E, 1984, p. 220. There are now revisions of Aphis for Britain
(Stroyan, 1984) and Fennoscandia/Denmark (Heie, 1986), and Brown (1989)
has keyed the alatae of the northwestern European species. A key to Aphis
of Manitoba, Canada, is also available (Rojanavongse and Robinson, 1977).

Aphis acanthopanaci Matsumura Apterae 2.0-2.3 mm, dark green accord-

ing to original description of specimens from Acanthopanax ricinifolium
(Matsumura, 1917); midsummer specimens (from Aralia cordata) about
1.3 mm and yellow according to Takahashi (1966). Only known from Japan.
Alatae undescribed, life cycle unknown.

Aphis arbuti Ferrari Apterae 1.7-2.7 mm, wine-red, on Arbutus unedo in

southern Europe. A member of the taxonomically difficult A. fabae group,
and only recognizable as a distinct species by its colour and host association.
Mier Durante and Nieto Nafria (1986) provide morphometric descriptions of
558 Aphis

apterae and alatae. Apparently monoecious and holocyclic (Barbagallo and

Stroyan, 1982), but sexual morphs are still undescribed.

Aphis asclepiadis Fitch Apterae yellowish-green, on Apocynaceae (Apo-

cynum) and Asclepiadaceae (Asclepias) in North America east of the Rocky
Mountains. Life cycle unknown (Rojanavongse and Robinson, 1977). 2n = 8.

Aphis caliginosa Hottes and Frison Small, brownish aphids with dark
legs and antennae, in ant-attended colonies on ends of branches of Cornus
racemosa in eastern North America (Illinois, Pennsylvania, North Carolina,
New Jersey). Life cycle unknown (Robinson and Chen, 1969).

Aphis catalpae Mamontova Apterae rich yellow to yellow-green, with a

dark green spot on dorsal abdomen in life. On undersides of leaves causing
strong wrinkling, and along petioles of Catalpa spp. (bignonioides, speciosa)
in Ukraine. Life cycle apparently not studied, but reported to migrate facul-
tatively to herbs and grasses (Mamontova, 1953).

Aphis celastrii Matsumura Apterae yellow or green with dark legs (but dark
greenish-brown according to original description), recorded from Celastrus
orbiculatus, Deutzia sp., Ilex serrata and Polygonum spp. in Japan (Higuchi
and Miyazaki, 1969). Closely related to A. spiraecola. Life cycle unstudied,
but oviparae have been collected on Deutzia (Takahashi, 1966). 2n = 8.

Aphis chetansapa Hottes and Frison Apterae yellowish-brown to yellowish-

red with darker antennae and legs and blackish siphunculi, and dark markings
on posterior abdominal tergites. Alatae have only 4-8 secondary rhinaria
confined to the third antennal segment, dark wing veins and often have dark
tranverse bands across last three abdominal tergites (Hottes and Frison, 1931).
In tightly curled terminal leaves of Pyrus sp. and Prunus sp., recorded from
Illinois, Missouri and Oregon, USA. The description of this species closely
resembles that of A. longicauda. Life cycle is unknown.

Aphis commensalis Stroyan Apterae 1.4-1.7 mm, dark grey-green to

blackish-green, heavily clothed in mealy greyish wax powder, living within
empty leaf-edge pocket-galls of the psyllid Trichopsylla walkeri on leaves of
Rhamnus cathartica in Europe. Small apterous males and oviparae mature
within the gall in September-October and eggs are laid within the gall
(Stroyan, 1952; or for a fuller account Böhm, 1963).

Aphis coreopsidis (Thomas) Apterae 1.5-1.8 mm, yellow to green with

darker antennae, legs and siphunculi, on stems and leaves of new shoots of
Nyssa sylvatica in spring. Apparently host-alternating in North America
(Illinois) between Nyssa and secondary hosts in Compositae (Bidens, Cliba-
dium, Eupatorium, Sonchus) or Malvaceae (Hibiscus, Sida). Oviparae in
autumn on Nyssa are small (1.1-1.2 mm), yellowish-green; males are alate,
about 1 mm, dark green and brown (Hottes and Frison, 1931). The fundatrices
 Aphis 559

on the primary host are still undescribed. Presumed anholocyclic populations

on Compositae (especially Bidens pilosa) occur in Central and South America,
and alatae have been trapped in Ghana.

Aphis cornifoliae Fitch Apterae are small (1.0-1.4 mm), dark brown to dull
greenish-black, on both upper and undersides of leaves of Cornus spp., living
without host alternation; males are apterous (Palmer, 1952). Widely distri-
buted in USA and Canada. Robinson and Chen (1969) discussed differences
from A. helianthi.

Aphis craccivora Koch Apterae 1.4-2.0 mm, dark brown to shining black,
without any wax dusting when adult, on young growth and usually ant-
attended. Polyphagous, but with a marked preference for Leguminosae.
Cosmopolitan. (See B & E, 1984, p. 223.) 2n = 8.

Aphis crinosa (Paik) Apterae yellowish-brown with white wax secretion; BL

c. 2.3 mm (Shinji, 1923, as Pterocomma ligustri). On trunks and branches of
Ligustrum spp. in Japan and Korea (Paik, 1965, 1972).

Aphis cytisorum Hartig Apterae 1.5-2.8 mm, very dark green to black and
covered with grey wax powder, ant-attended on young shoots and petioles
and later on inflorescences of Laburnum anagyroides; also recorded from
Spartium junceum. Monoecious holocyclic, with alate males. Widely distri-
buted in Europe eastward to Bulgaria and Russia; introduced and widespread
in North America. See Stroyan (1984), Heie (1986). 2n = 8.

Aphis eugeniae van der Goot Apterae 1.4-1.6 mm, orange-yellow to

brownish-orange with black siphunculi and CAUDA. Resembling A. spiraecola,
but distinguishable by the characters given in the key to aphids on Glochidion.
Most commonly on Euphorbiaceae (Glochidion, Breynia, Phyllanthus), but
plants in other families may be colonized (Eugenia, Dipsacus, Chromolaena).
Throughout East and Southeast Asia, and in Australia (Eastop, 1966, as
A. hardyi). Calilung (1976) gave an account of this species in the Philippines.
2n = 8.

Aphis euonymi Fabricius Apterae various shades of chocolate brown, often

with transverse bars of wax dust on anterior abdominal tergites; otherwise
similar to A. fabae. Monoecious and holocyclic on Euonymus europaeus,
curling the leaves. Males, and usually the gynoparae also, are apterous.
Hybridization with A. fabae occurs readily in the laboratory and may occur
in the field, so subspecies status might be more appropriate for this aphid
(Müller and Steiner, 1986). Heie (1986) provided a general account, and
discussed the confused application of this name in the literature. 2n = 8.

Aphis fabae Scopoli Apterae 1.5-3.1 mm, dull black, sometimes with white
wax markings, usually ant-attended. Curling leaves of Euonymus europaeus
(also Philadelphus coronarius and Viburnum opulus) in spring and migrating
560 Aphis

to a wide range of secondary hosts, including some trees (e.g. Aesculus,

Catalpa, Prunus, and even Larix). Oviparae on Euonymus in autumn are
small, with strongly swollen hind tibiae, and males are alate (Fig. 4). In reality
a complex of sibling species or partially interfertile host races, of which four
are known to overwinter on Euonymus; they are fabae sensu stricto (which
in host transfers will colonize Vicia faba), cirsiiacanthoides Scopoli (which
will transfer to Cirsium arvense), solanella Theobald (which will transfer to
Solanum nigrum) and euonymi, dealt with separately above. Accounts of this
group in Europe were given by Stroyan (1984) and Heie (1986), and the most
recent taxonomie treatment was by Müller (1988). Outside Europe, anholo-
cyclic populations of A. fabae group on secondary hosts occur in Southeast
Asia, Africa, Indian subcontinent, South America, Hawaii and Auckland
Isles. Warm temperate and subtropical populations seem to correspond best
to solanella (B & E, 1984, p. 224). 2n = 8.

Aphis farinosa Gmelin Apterae 1.6-2.5 mm, dull green with CAUDA dis-
tinctly darker than the long pale siphunculi, in dense colonies on young shoots
of Salix spp. in spring and early summer, attended by ants. Monoecious, with
green oviparae and orange males appearing from July onwards, although
occasionally populations of viviparae persist until August or September
(Stroyan, 1984; Heie, 1986). Occurs throughout northern temperate parts
of the world, and in South America (Argentina). A nominal subspecies,
yamagicola Matsumura, occurs commonly on Salix spp. (daphoides, rorida)
in Japan and is distinguished by having long, fine hairs on the third antennal
segment (about twice the middle diameter of the segment); it produces sexuales
in the autumn (Takahashi, 1966). Populations on Salix gracilistyla in
Siberia have short antennal hairs like European farinosa, but a longer last
rostral segment; R IV+V 1.27-1.45 x HT II, as compared with 1.07-1.25 x
HT II (Holman, 1987). 2n = 6.

Aphis feminea Hottes Apterae red with black head, pro- and mesothorax,
antennae, legs, siphunculi and CAUDA, lightly dusted with white wax. It
forms dense colonies on young twigs and small branches of Prunus serotina in
northeastern USA (Hottes and Frison, 1931).

Aphis frangulae Kaltenbach Apterae 1.4-2.4 mm, yellow, green, brown or

black, curling leaves of Rhamnus frangula (= Frangula alnus) in spring. The
name is applied to a complex of sibling species or host races utilizing Rhamnus
as primary host in Europe (frangulae, beccabungae, testacea), populations of
which may show colour differences in life (see Heie, 1986). (The complex
also includes A. gossypii which is anholocyclic in Europe - see below.) The
taxonomie problems of the frangulae group were discussed by Stroyan (1984).
Of the nominal subspecies forming spring populations on Rh. frangulae
in continental Europe, frangulae sensu stricto will preferentially colonize
Epilobium angustifolium and beccabungae will colonize Veronica beccabunga,
Solanum tuberosum and various Labiatae, whereas testacea (only known
from Germany) seems to be monoecious on Rhamnus. However, natural
 Aphis 561

hybridization may tend to confuse secondary host associations and life cycle
categories, and there are also probably geographical differences. 2n = 8.
Aphis utilis Zhang is a closely-related species in China.

Aphis glycines Matsumura Host-alternating between Rhamnus dahurica

and Glycine max in China (Wang et al., 1962); elsewhere in East and Southeast
Asia only recorded from Leguminosae (Glycine, Pueraria, Desmodium).
Appearance of colonies in life on Rhamnus unrecorded; aphids on secondary
hosts are small and yellow with black siphunculi (B & E, 1984, p. 225).

Aphis gossypii Glover In Europe A. gossypii is treated as an anholocyclic

and polyphagous member of the A. frangulae group (see above), and anholo-
cyclic populations usually identified as gossypii occur throughout the world
on many different plants, sometimes including tree species. However, Kring
(1959) demonstrated a facultative holocycle in Connecticut, USA, utilizing
Catalpa bignonioides and Hibiscus syriacus as primary hosts, and a holocycle
is common in Japan (Takada, 1988) and China (Zhang and Zhong, 1982a).
Primary hosts in East Asia include Celastrus orbiculatus, Hibiscus syriacus,
Rhamnus dahurica, Rh. japonica and Punica granatum. Zhang and Zhong
also record a monoecious holocycle for gossypii on both Hibiscus and cotton.
The taxonomy of populations under the name gossypii in different parts of
the world is therefore extremely confused. Apterae of holocyclic populations
on primary hosts are of varying colour, usually greenish, but aphids in summer
on woody plants are usually small and yellowish. Males are always alate
(B & E, 1984, p. 226). 2n = 8.

Aphis hederae Kaltenbach Apterae dark brown; BL 1.4-2.5 mm. Immatures

paler brown. Widespread in north temperate regions on young shoots and
leaves of Hedera helix, and frequently on other Araliaceae including Schleffera
spp. growing in glasshouses and conservatories. 2n = 8.

Aphis helianthi Monell Host-alternating between Cornus stolonifera and

various herbaceous plants, particularly Umbelliferae. Spring populations twist
and curl the leaves of Cornus (Hottes and Frison, 1931); apterae are 1.5-
1.8 mm, dark yellow to green, with siphunculi variably pigmented from green
to black. Oviparae in autumn have hind tibiae considerably swollen, and males
are alate. Widespread in North America and also recorded from Brazil on
Yucca (assuming A. yuccicola Wilson is a synonym). (See also Robinson and
Chen, 1969.) 2n = 8.

Aphis horii Takahashi Apterae blackish-green, clustered on stems and leaf

petioles of Sambucus spp. in Japan. Paik (1965) also recorded this aphid (as
Sappaphis euscaphis) in Korea from Euscaphis japonica (Staphyleaceae).
Closely related to A. sambuci, but differing in the presence of long, fine hairs
on the head and antennae. Host alternation to roots of Caryophyllaceae, Poly-
gonaceae, etc., as in A. sambuci, is not recorded in the literature, but speci-
mens have been collected in Korea on Rumex coreanus by W.H. Paik (BMNH
562 Aphis

colln). Aphis horii was originally described from Cirsium dipsacalepsis

(Takahashi, 1923), but Takahashi (1966) regards this host record as doubtful.

Aphis ilicis Kaltenbach Apterae 1.7-2.9 mm, dark olive-brown, reddish or

greyish-brown, living in dense colonies on young shoots and undersides of
young leaves of Ilex aquifolium. Attacked leaves curl towards their under-
sides. Immatures often have white pleural wax markings (it is a member of
the A. fabae group). Apparently monoecious, although it is unclear how
colonies survive through the summer under natural conditions, as mature
leaves are not colonized. Sexuales are recorded from July onwards; males are
alate. Occurs in most of Europe, eastward to Turkey. North American records
require confirmation. (See also Stroyan, 1984; Heie, 1986.) 2n = 8.

Aphis kachkouli Remaudière Apterae pruinose; in spring 1.2-1.5 mm and

pale green, in autumn only 0.5-0.8 mm and pale yellow or whitish. In some-
times dense colonies on leaves of Ficus carica, and F. johannis (probably the
original host) in Iran (Remaudière, 1989). Monoecious, with apterous males.
Antennae are 5-segmented in both apterae and alatae, and also in males
(sometimes 4-segmented in dwarf apterae).

Aphis kalopanacis (Hori) Apterae about 1.9-2.0 mm, purplish-black, on

undersides of leaves and petioles of Kalopanax ricinifolius in Hokkaido,
Japan (Hori, 1927). Monoecious, with alate males. Apparently not recorded
since the original description (as Pergandeida kalopanacis).

Aphis lantanae Koch (= setacea Hille Ris Lambers) Apterae about 1.6mm,
colour in life probably black, resembling A. fabae. In Western Europe,
normally monoecious on Viburnum lantana (see Stroyan, 1984). Specimens
collected on Sorbus aria in Bavaria, Germany, in 1933 were described as new
under the name Doralis setacea, but this name was synonymized with A.
lantanae by Eastop and Hille Ris Lambers (1976).

Aphis longicauda (Baker) Apterae about 1.7-1.8 mm, of variable colour,

brown or yellowish-white with dark antennae and legs and dark posterior
abdominal markings. On twigs and leaves of plum trees (Prunus domestica) in
USA, recorded as very injurious to plum in Virginia, with twigs dying the year
after attack (Baker, 1920), but little known since original description. Monoe-
cious, with apterous males. The description resembles that of A. chetansapa.

Aphis maculatae Oestlund Host-alternating between Cornus stolonifera and

Populus spp., where it sometimes forms large colonies on twigs and leaves,
throughout most of North America. Apterae on Populus in summer are
brownish-black, with black siphunculi and CAUDA and paler head, legs and
antennae, the body marked with conspicuous patches of white wax powder
(Palmer, 1952). Wilson and Moore (1986) studied the differential susceptibility
of various Populus hybrids to this species and discussed insecticidal control
measures.
 Aphis 563

Aphis madronae (Essig) Apterae about 2.1-2.2 mm, reddish-brown through-

out, in small or large dense colonies on trunk and lower leaves of Arbutus
menziesi in California. The aphids match the colour of the bark of the tree.
Not recorded on Arbutus since the original collection, but specimens from
Arctostaphylos sp. (BMNH colln, leg. R.C. Dickson) are identified as this
species. Life cycle is unknown.

Aphis mammulata Gimingham and Hille Ris Lambers Apterae 1.3-1.9 mm,
greenish-grey, found in autumn in dense, ant-attended colonies on petioles of
berries and undersides of leaves of Rhamnus cathartica. Monoecious holo-
cyclic; the colonies include oviparae and orange-brown males. The alata is
described by Heie (1986). Appearance of the colonies earlier in the season is
unrecorded. Recorded from England, Sweden, Germany, Poland, Czechoslo-
vakia and Russia (as A. rhamnicola Mamontova).

Aphis minima Tissot Apterae about 1.0-1.1 mm, yellowish-brown to dark

reddish-brown with darker siphunculi and CAUDA, on young twigs of Prunus
americana and also feeding along main veins of leaves, which may become
tightly curled. Only recorded from Florida, USA. Monoecious holocyclic,
with apterous males (Tissot, 1933).

Aphis nasturtii Kaltenbach Host-alternating between Rhamnus cathartica

(and Rh. alnifolia) and a wide variety of secondary hosts (B & E, 1984, p. 230).
Apterae on Rhamnus in spring are rather bright green to greenish-yellow and
cause distortion of young leaves. Widely distributed in Europe and across Asia
to Japan; introduced into and widespread in North America. (See also Heie,
1986.) 2n = 8.

Aphis neogillettei Palmer Apterae 1.0-1.4 mm, dark olive-green with

blackish siphunculi, CAUDA and apices of appendages, in dense colonies
curling leaves of Cornus stolonifera. Monoecious holocyclic, with olive-
brown apterous males (Palmer, 1952). Widely distributed in North America.
2n = 8.

Aphis nerii Boyer de Fonscolombe Apterae 1.5-2.6 mm, bright yellow with
black siphunculi and CAUDA; antennae and legs also predominantly dark.
Alatae have dark wing veins. Mainly colonizing Asclepiadaceae and Apo-
cynaceae, often in large, dense colonies on young stems, and occasionally
on plants (including trees) of other families (e.g. Catalpa, Citrus). Widely
distributed throughout the tropics including many Pacific islands. Apparently
mainly anholocyclic, but sexual morphs were recently described from Japan
(Takada and Miyazaki, 1992). (See also B & E, 1984, p. 230; Stroyan, 1984.)
2n = 8.

Aphis nigra Wilson Apterae about 0.8mm, greyish-black with yellowish

antennal flagellum and tibiae. Collected on Cyrilla racemiflora at Batesburg,
South Carolina, USA, in 1910, and not since recorded (Wilson, 1911).
564 Aphis

Aphis nigratibialis Robinson Apterae 1.3-2.1 mm, blackish in colour with

tibiae also wholly black. On Cornus stolonifera in Idaho, Oregon and Utah,
USA. Life cycle is unknown (Robinson and Chen, 1969).

Aphis pawneepae Hottes Apterae 1.5-1.7 mm, brown to reddish-brown

with antennae and legs wholly dusky brown, found by Hottes (1934) in shelters
constructed by the ant Crematogaster lineolata at the bases of trunks of young
Cercis canadensis trees. Parker (1935), however, found colonies arranged
along undersides of twigs and branches (always on old wood). Fundatrices and
oviparae have also been collected, so this species is monoecious holocyclic
on Cercis. Recorded from Illinois, Kansas, Montana, North Carolina and
Pennsylvania, USA.

Aphis pomi De Geer Apterae 1.3-2.2 mm, bright apple-green with black
siphunculi and dark CAUDA, in dense colonies on young growth of several
genera of woody Rosaceae (Pomoidea); ant-attended, and causing slight leaf
curl (B & E, 1984, p. 231). Later generations occur on the undersides of leaves
and are yellow-green. Monoecious holocyclic, with apterous males. In Europe,
southwest Asia and North America (where there is some confusion in the
literature with A. spiraecola on the same hosts). Alatae have darker wing veins
than A. spiraecola; Halbert and Voegtlin (1992) gave some other discriminants
based on North American populations of the two species. Hogmire et al.
(1992) compared population development and insecticidal susceptibility of
A. pomi and A. spiraecola. Oriental records of A. pomi all seem to apply
to other species, although it is surprising that it has not spread worldwide. (See
also Heie, 1986.) 2n = 8.

Aphis ramnifila David, Narayanan and Rajasingh Apterae 1.1-1.8 mm,

blackish-green, with brown legs and antennae, and darker siphunculi and
CAUDA, collected on undersides of leaves of Rhamnus virgatus in India
(Uttar Pradesh, Himachal Pradesh, Garhwal Himalaya). Possibly this name
is being applied to more than one species (David et al., 197la).

Aphis rhoicola Hille Ris Lambers Apterae 1.7-2.1mm, brown with pale
antennae and legs (except black tarsi) and black siphunculi. On leaves and
branches of Rhus abyssinica in Eritrea (Hille Ris Lambers, 1954a). Resembling
A. gossypii, but with a consistently shorter antennal terminal process. Life
cycle is unknown; however, very similar aphids occur on Rumex in India
(BMNH colln), so this is a possible secondary host.

Aphis roumanica Holman Apterae 1.4-1.9 mm, green; appearance in life

unrecorded but probably similar to the closely related A. farinosa. Monoe-
cious holocyclic on Salix purpurea in Roumania, with oviparae collected in
July (Holman, 1990). Alate viviparae and males undescribed.

Aphis salicariae Koch (= corniella Hille Ris Lambers) Apterae 1.6-2.4 mm,
reddish-brown with a light dusting of wax; legs pale (except tarsi), siphunculi
 Aphis 565

and CAUDA black. Causing leaf curl to Cornus spp. (alba, sanguinea,
stolonifera) in spring and also feeding in developing flower umbels; host-
alternating to Epilobium angustifolium. Often ant-attended. Widely distri-
buted in Europe (except Spain and Portugal), eastward to Central Asia; and
introduced into and widespread in North America. 2n = 8.

Aphis sambuci Linnaeus Apterae 1.9-3.5 mm, dark green or dark brown to
yellowish-brown, in dense colonies around young stems of Sambucus spp. in
spring. Host alternation occurs to roots of various plants, especially Caryo-
phyllaceae and Rumex (Stroyan, 1984; Heie, 1986). Colonies may persist
through summer on Sambucus and may produce oviparae, but males only
seem to be produced on secondary hosts. In mild winters in England apterae
may hibernate on subterranean parts of Sambucus. Occurs throughout Europe
and temperate Asia (except that in Japan it seems to be replaced by a long-
haired relative, A. horii) and North and South America. It is often recorded
in North American literature as sambucifoliae Fitch. 2n = 8.

Aphis sassceri Wilson Apterae about 1 mm, greenish-brown with dusky

siphunculi; alatae predominantly orange-yellow. Apterae have distinct mar-
ginal abdominal tubercles. Known only from the original collection on
Annona 'rectilineata' (= ? reticulata) in Florida, USA (Wilson, 1911).

Aphis schinifoliae Blanchard Apterae 1.4-1.7 mm, blackish-brown in life

with antennae and legs mainly dark and CAUDA dark, but siphunculi pale,
rather short and almost flangeless. The ANT PT is short (ANT PT/BASE
ratio about 1.4) and there are well-developed abdominal marginal tubercles
(Blanchard, 1939). On Schinus spp. (dependens, molle) in Argentina and
Chile. Alata undescribed and life cycle is unknown.

Aphis sophoricola Zhang Apterae blackish with whitish wax bloom, col-
lected on Sophora japonica at Beijing, China (Zhang and Zhong, 1981a).
Closely related to, if not synonymous with, A. cytisorum.

Aphis spiraecola Patch (= citricola van der Goot of recent authors) Apterae
1.2-2.2 mm, bright greenish-yellow to apple green, with brownish head and
blackish siphunculi and CAUDA; in colonies often distorting growing points,
usually ant-attended. Very polyphagous, particularly on plants of shrubby
habit, but also often on trees. Anholocyclic through most of the world, but
a holocycle involving Spiraea spp. as primary hosts is recorded from North
America, Brazil and Japan; in the latter country, Citrus is also a primary host
(B & E, 1984, p. 222, as A. citricola). 2n = 8.

Aphis taraxacicola (Nevsky) Apterae about 1.7-1.8 mm with pale yellow

body and appendages and short siphunculi (only about 0.1 x BL, just a little
longer than CAUDA). Alatae have a pale yellow abdomen with brown pig-
ment medially on tergites 1-6 and only about two secondary rhinaria on the
third antennal segment. This species is described both from undersides of
566 Aploneura

leaves of Aesculus hippocastanum in June, and from inside leaf rosettes of

Taraxacum spp. in June and August, in Central Asia (Nevsky, 1951). It has
not since been recorded. Biology and life cycle are unknown.

Aphis utilis Zhang Apterae about 1.4mm, appearance in life unrecorded.

Collected on Rhamnus utilis at Beijing, China (Zhang and Zhong, 1983a).
Apparently a member of the A. frangulae group.

Aphis verbasci Schrank Apterae 1.7-2.5 mm, bright golden yellow to pale
green with black siphunculi, ant-attended, on undersides of leaves of Buddleja
spp. (farreri, japonica, madagascariensis, paniculata), Verbascum spp., and
also recorded from Scrophularia. Monoecious holocyclic on these hosts;
sexuales are undescribed, but there are oviparae and apterous males in the
BMNH collection. In Europe (except Fennoscandia), Middle East, North
Africa, India and Pakistan. 2n = 8.

Aphis versicolor (Börner) Apterae 1.1-1.6 mm, body and siphunculi dark
green, on leaves of Rhamnus cathartica in Austria and Bulgaria (Börner,
1950). This species has similar very large marginal tubercles to A. mammulata
on this host from other parts of Europe, but much shorter hairs on antennae
and marginal areas of abdomen. Possibly it should be regarded as a geo-
graphical variant of that species. Monoecious holocyclic; males are apterous
and orange-brown, as in mammulata.

Aphis viridissima Mamontova Apterae 1.6-1.7 mm, bright green, on ter-

minal shoots of Tamarix sp. in Ukraine (Mamontova, 1955). Other morphs
and life cycle are unknown.

Aphis viticis Ferrari Apterae 0.8-1.7 mm, grass green with brown siphunculi
in spring, pale yellow in midsummer. Monecious holocyclic on Vitex agnus-
castus, with alate males; recorded from France, Iran, Israel, Italy, Mallorca,
Spain and Turkey. Barbagallo and Stroyan (1982) redescribed the apterous
and alate viviparae, and Nieto Nafria et al. (1986) described the sexuales.

Aphis whiteshellensis Rojanavongse and Robinson Apterae 1.5-1.8 mm,

reddish-brown, in tightly curled leaves at ends of small branches of Amelan-
chier alnifolia in Manitoba, Canada; also collected from Amelanchier sp. in
Montana, USA (BMNH colln, leg. D.J. Voegtlin). Monoecious holocyclic,
with apterous males (Rojanavongse and Robinson, 1977).

APLONEURA Passerini Pemphiginae: Fordini

A small genus with little to distinguish it morphologically from Slavum.

However, the alatae of Aploneura rest their wings flat over the abdomen,
 Apulicallis 567

whereas in Slavum the wings are held roof-like. The life cycle of only one
species is known and that has a specific primary host association with Pistacia
lentiscus.

Aploneura lentisci (Passerini) Host-alternating between Pistacia lentiscus

and the roots of Gramineae, with the two-year life cycle that is typical of For-
dini, throughout the Mediterranean region. Anholocyclic populations on grass
roots have spread to most other parts of the world except North America
(B & E, 1984, p. 235). The galls are pocket-like, often kidney-shaped, formed
by invagination of the leaf alongside the mid-rib (Fig. 126A). They appear in
late March-April (in Israel) and grow quickly to attain their maximum size
by May, although the fundatrix inside does not start to reproduce until early
June. Galls open in August-November and emigrant alatae (1.3-2.3 mm) leave
as they become mature, over an extended period. Wool and Manheim (1986,
1988) studied the population ecology of gall generations in Israel. 2n = 16.

APPENDISETA Richards Drepanosiphinae: Phyllaphidini

A genus with one North American species related to Pterocallis, but with two
pairs of anterior prothoracic marginal hairs and a small hair arising from the
siphunculus, on the ventral side near its base (Richards, 1965).

Appendiseta robiniae (Gillette), Plate 9a Viviparae all alate, pale yellow-

green with spinopleural and marginal longitudinal rows of pale powdery spots,
on undersides of leaves of Robinia pseudacacia and R. neomexicana, and also
now recorded from Sophora japonica (Forbes and Chan, 1989). Sexuales
(alate males and apterous oviparae) occur in September-November (Palmer,
1952). Widespread in North America, and introduced into Europe (England,
Germany, Hungary, Italy, Spain, Switzerland) and the Middle East (Jordan:
BMNH colln). 2n = 10*.

APULICALLIS Barbagallo and Patti

Drepanosiphinae: Phyllaphidini

A genus for one species in Italy related to Myzocallis, but with one pair of
spinal hairs on each abdominal tergite in all morphs and with other differences
in larval chaetotaxy.

Apulicallis trojanae Barbagallo and Patti Alatae lemon yellow with head
and thorax very slightly brownish; BL 1.3-2.3 mm. Holocyclic and mono-
568 Asiphonaphis

phagous on Quercus trojana in southeast Italy. Oviparae and alate males

in November. Barbagallo and Patti (1991) described all morphs.

ASIPHONAPHIS Wilson and Davis Aphidinae: Aphidini

A small North American genus with only two currently recognized species,
related to Aphis but completely lacking siphunculi. Both alatae and apterae
have prominent marginal tubercles on all abdominal segments.

Asiphonaphis pruni Wilson and Davis Apterae are pale yellow to whitish-
green, sometimes banded with darker green, with darker head and pronotum,
in colonies on leaves and distorting the new growth of Prunus spp. (armeniaca,
serotina, virginiana) in Canada and USA. Monoecious holocyclic, with
apterous males. Robinson (1964) gives an account of this species on P. virgi-
niana in Manitoba.

ASIPHONELLA Theobald Pemphiginae: Fordini

The two species of this genus form large, cockscomb-like galls on Pistacia and
migrate to roots of grasses, especially Cynodon dactylon, with a holocycle
taking two years. Accounts are given by Remaudière and Tao (1957) and
A.K. Ghosh (1984b).

Asiphonella cynodonti (Das) Host-alternating between Pistacia khinjuk and

roots or ground level parts of Cynodon dactylon in Iran, Pakistan and
India, and also found galling P. palaestina in Israel (Wool, 1984). Galls are
formed by the invagination of the leaf alongside the mid-rib to form a large,
cockscomb-like structure (Fig. 126B). The emigrant alatae (BL about 1.8-
1.9 mm) leave through secondary openings in the wall of the gall in September
to November.

Asiphonella dactylonii Theobald Host-alternating in China between Pistacia

sinensis and roots and ground level parts of Cynodon dactylon and certain
other grasses, with anholocyclic populations on grass roots in other parts of
the world (B & E, 1984, p. 237; see also Silva, 1985). Galls are similar to those
of A. cynodonti, but mature earlier in the year with alate emigrants (BL
1.5-1.9 mm) collected in May (Remaudière and Tao, 1957).
 Asíegopteryx 569

ASPIDOPHORODON Verma Aphidinae: Myzini

A small genus associated with Salix in East Asia. The antennae are 4- or
5-segmented with terminal process 0.6-1.0 times base of last segment and
the siphimculi have clavate ends with the aperture small and subapical.
The head of the aptera bears anteriorly-directed medial and lateral frontal
projections.

Aspidophorodon harvensis Verma Apterae of BL 1.5-1.9 mm, mainly pale

yellowish with pale brown head; alatae with black head and thorax and
brownish transverse bars on dorsal abdomen. On undersides of leaves of an
unidentified Salix species, sitting tightly alongside the veins, in Kashmir, India
(Verma, 1965). Life cycle is unknown.

Aspidophorodon salicis Miyazaki Apterae are whitish, pale yellow or pale

yellowish-green, BL 1.4-1.6 mm in spring; dwarf summer apterae have BL
0.9-1.4 mm and 4-segmented antennae. Alatae have a black head and thorax
and a large dark central abdominal patch. On undersides of leaves of Salix
sachalinensis in Hokkaido, Japan (Miyazaki, 1971), and also recorded from
Sakhalin and the Kuril islands (Pashchenko, 1988b). Aspidophorodon sini-
salicis Zhang, described from Salix sp. in China (Zhang and Zhong, 1980a),
is closely related and possibly the same species.

ASTEGOPTERYX Karsh Hormaphidinae: Cerataphidini

An oriental genus of about 30 species with host alternation between Styrax

and monocotyledonous plants, chiefly palms and bamboos. The galls on
Styrax vary greatly in morphology between species and are sometimes very
elaborate (Fig. 124). The morphs from these galls differ greatly from those on
secondary host plants, which were described in different genera (Trichoregma,
Oregma) until Hille Ris Lambers (1953b) showed that Astegopteryx styra-
cophila on Styrax benzoin was the same species as Oregma pallida Van
der Goot on bamboos. Apterae of Astegopteryx on bamboos and palms are
small aphids with well-developed frontal horns and segmentally arranged wax
glands, forming large colonies on the leaves. Most species probably have the
ability to maintain themselves continuously by parthenogenetic reproduction
on their secondary host plants. The bamboo feeders seem to fall mainly into
three groups, exemplified by A. bambusae, A. basalis and A. styracophila.
Noordam (1991) reviewed the Javanese species. For palm-feeding species see
B & E, 1984, pp. 238-239.
570 Asíegopteryx

Astegopteryx bambusae (Buckton) Broadly pear-shaped aphids, BL 1.2-

2.0mm, often distinctively coloured in life, with rounded apices to their
frontal horns, generally colonizing the undersides of the leaves of Bam-
busa and other bamboos. Alatae occur (in Java) from September to January
(Noordam, 1991). Members of the bambusae group occur in the Indian
subcontinent and throughout East and Southeast Asia. Several closely-
related species have been described mainly on the basis of differences in
colour and appearance in life, although some are based on differences in
antennal morphology and wax gland development seen in mounted pre-
parations. Migration to Styrax is known for only one species in this group,
A. bambucifoliae (q.v); probably other species either live exclusively on
bamboo or have primary host generations known by other names on Styrax.
Although populations in any one locality at one time may show consistent
colour and wax gland development, the extent to which climate and other
environmental factors influence these characters is uncertain, and detailed
study, including rearing work, is necessary to make sense of the morphological
variation in this group when populations from more than one country are
considered.
Astegopteryx bambusae was originally described from Uttar Pradesh,
India, colonizing the upper sides of leaves of Bambusa arundinacea (Buckton,
1893). Buckton recorded the colour as greenish-brown, mottled with black, but
he was presumably looking at dead specimens. Doncaster (1966) redescribed
the Buckton material and made Oregma lutescens van der Goot from Java
a synonym. According to van der Goot's original description, lutescens is
yellow-brown with indistinct dorsal transverse bands on abdominal tergite 1
and between the siphunculi. Noordam (1991) placed similis van der Goot, 1917
and striata van der Goot, 1917 as additional synonyms of bambusae, regarding
them, with lutescens, as colour varieties (see Noordam, 1991, pp. 1-5). He also
considered insularis van der Goot (1912, nee 1917) to be a synonym of bam-
busae. 2n = 12 (Kar et al., 1990).

Astegopteryx bambucifoliae (Takahashi) Host-alternates between Styrax

suberifolium and various bamboos in Taiwan. The generations galling Styrax
were described as Astegopteryx sasakii (Takahashi, 1936b), but were linked
with the bamboo-feeding generations by Aoki and Kurosu (1989). The galls
comprise about 20 elongate subgalls radiating from one point on the stem
(Fig. 124A). Kurosu and Aoki (1990a) reported 'chimeral' galls formed initially
by A. bambucifoliae, but transformed by invading Ceratoglyphina styracicola.
Alatae (BL about 1.7mm) emerge from distal openings in the subgalls in
August-November to fly to bamboos (Kurosu and Aoki, 1991b). The secon-
dary host generations form spaced-out colonies on undersides of leaves of
Bambusa spp. (dolichoclada, edulis, oldhami, stenostachya), Dendrocalamus
latiflorus and Phyllostachys lithophia. Apterae are yellowish-green, with two
darker green longitudinal stripes and wax tufts on the sides of the abdomen;
BL 1.4-2.1mm (Liao, 1976). Defensive behaviour patterns involving use
of the frontal horns for butting are described by Aoki and Kurosu (1985,
1989).
 Astegopteryx 571

Astegopteryx basalis van der Goot The name applied to bamboo-feeding

populations in Southeast Asia, which are similar to A. bambusae, but have
broadly conical pointed frontal horns. Astegopteryx basalis was described
from unidentified Bambusa species in Java, as a yellowish-white aphid, BL
1.1-1.7 mm, with paired brownish-red areas on the dorsum and with wax
secreted especially on the sides of the abdomen (van der Goot, 1917). Noordam
(1991) described other colour forms in Java and placed three Takahashi
species in synonymy: vandermeermohri (Takahashi, 1935a) from Sumatra;
flava (Takahashi, 1950), apterae of which are entirely yellow with paired
sclerotic patches on abdominal tergites 1-4 and transverse bars on 5-7; and
malaccensis (Takahashi, 1950), a dark purplish-brown form with very well-
developed wax glands from Malaysia.

Astegopteryx chinensis Tao Forms globular galls similar to those of Cera-

toglyphina bambusae on an unidentified species of Styrax in Szechuan, China
(Tao, 1966). Alatae (BL about 1.8mm) are mature in the galls in July, and
fly to an unknown secondary host.

Astegopteryx glandulosa Noordam Apterae are yellow, with orange mark-

ings or marbling, wholly covered with wax wool which extends as a wax
powder onto the leaf; BL 1.2-1.5 mm. Immatures are bright yellow, develop-
ing orange spots. On undersides of leaves of unidentified bamboos in Java,
at altitudes above 1100m (Noordam, 1991).

Astegopteryx insularis (van der Goot) Originally described from an uniden-

tified host plant in Java (van der Goot, 1912), but this material was syn-
onymized with A. bambusae by Noordam (1991). The material from bamboos
in Java later described under this name (van der Goot, 1917) was redescribed
by Noordam as A. unimaculata (q.v.).

Astegopteryx jamuritsu Takahashi Galls are large (12.0-14.5 cm long), pen-

dulous, flask-shaped, greyish-brown dusted with white powder, broadest at
the distal end where there is a very large, slit-like opening (Fig. 124B); on
Styrax suberifolium in Taiwan. Aphids emerge from the galls and drop when
disturbed, and may sting by piercing the skin of the intruder. Alatae (BL about
1.5 mm) are mature in the galls in November, flying to an unknown secondary
host (Takahashi, 193la, 1934b).

Astegopteryx leeuweni Takahashi Galls are formed in the flower-heads of

Styrax serrulatus var. mollissimus in Sumatra. They are slender, pod-like,
downy, 5-8 cm long, with a distal opening (Fig. 124D). Alatae, BL about
1.6mm, were collected from them in June (Takahashi, 1936b).

Astegopteryx liukueinensis Liao Apterae are yellow in life with a pair

of longitudinal green markings, BL 1.4-1.6 mm, and occur on Bambusa
multiplex in Taiwan (Liao, 1976).
572 Astegopteryx
 Astegopteryx 573

Fig. 124. Galls of Cerataphidini on Styrax. A, Astegopteryx bambucifoliae on S. suberifolium (from

Takahashi, 1939b); B, A. jamaritsu on S. suberifolium (from Takahashi, 1934b); C, Cerataphis
?fransseni on S. benzoin (from Takahashi, 1936b; as Astegopteryx lambersi); D, A. leeuweni on Styrax
sp. (from Docters van Leeuwen-Reijvaan and Docters van Leeuwen, 1926); E, A. roepkei on
S. sumatranus (from Docters van Leeuwen-Reijnvaan and Docters van Leeuwen, 1926);
F, A. sumatrana on Styrax sp. (from Docters van Leeuwen-Reijnvaan and Docters van Leeuwen, 1926);
G, Tuberaphis taiwana on S. formosana (from Takahashi, 1934b, as Astegopteryx); H, Astegopteryx
vandermeermohri on Styrax sp. (after Docters van Leeuwen-Reijnvaan and Docters van Leeuwen, 1926);
I, Tuberaphis takenouchii on S. japonica (after Aoki and Usuba, 1989, as Astegopteryx);
J, Ceratoglyphina bambusae on S. suberifolium (from Takahashi, 1934b); K, Ceratovacuna nekoashi on
S. suberifolium (from Kurosu and Aoki, 1990b); L, Pseudoregma koshunensis on S. suberifolium (from
Takahashi, 1939b, as Astegopteryx shitosanensis).
574 Astegopteryx

Astegopteryx minuta (van der Goot) The name most commonly applied to
populations of bambusae-like aphids with marginal wax pore plates reduced
(so that they are small, circular and discontiguous) or totally absent. In life
such apterae are generally yellowish-white to pale green with two irregular
dark green longitudinal markings and scanty wax; BL 1.1-1.8 mm (Noordam,
1991). On undersides of leaves of bamboos (Bambusa, Dendrocalamus).
Populations in which the apterae have no visible marginal wax pore plates and
are not clearly distinguishable from A. minuta have been described from India
as new species: A. neelagiriensis David, 1959a, on Bambusa sp. in Madras;
and Pseudastegopteryx himalayensis M.R. Ghosh, Pal and Raychaudhuri,
1977b, on unidentified bamboo in West Bengal. Alatae occur (in Java) from
May to mid-October (Noordam, 1991). 2n = 12 (in India; Kar et al., 1990).

Astegopteryx pallida van der Goot Apterae greenish-white or yellow with

variable dark green dorsal markings and white wax, especially on margins; BL
1.1-2.2 mm (Noordam, 1991). On undersides of leaves of bamboos (Bambusa
spp.) in Java. Oregma salatigensis van der Goot (1917) is a synonym, and
possibly also Trichoregma bambusae Takahashi (1935a). It was suggested by
Hille Ris Lambers (1953b) that this aphid was the secondary host form of
A. styracophila, but this awaits experimental confirmation.

Astegopteryx pandani (Takahashi) Apterae pale brown to brown with

whitish antennae and legs and long marginal tufts of white wax; BL c. 1.4 mm.
On Pandanus tectorius in Sumatra (Takahashi, 1935a), and redescribed from
Freycinetia javanica in Java (Noordam, 1991). On undersides of leaves,
attended by ants. Other morphs and life cycle unknown.

Astegopteryx roepkei Hille Ris Lambers The gall of this species, found on
Styrax paralleloneurus in Sumatra, is remarkable; it is composed of thin,
spirally-twisted tubular subgalls, 15-20 cm long, their bases radiating from a
circular attachment site on one side of the stem near an axil-bud (Fig. 124E).
Alatae, BL about 1.8 mm, are adult in September. Secondary host is unknown.

Astegopteryx singaporensis (van der Goot) Apterae are yellowish-white or

pale brownish-yellow, with 1-3 pairs of small pale greenish spots, a somewhat
shiny, bright yellow to brownish area on and anterior to the siphunculi, and
variably developed marginal plumes of wax; BL 1.2-1.7 mm. On undersides
of old leaves of bamboos, which often show yellow or brown patches.
Originally described from unidentified bamboos in Singapore (van der Goot,
1918), and redescribed from Java, where it is recorded from species of
Bambusa, Dendrocalamus, Melocanna and Schizostachyum (Noordam,
1991). Alatae are unknown.

Astegopteryx styracophila Karsh Galls on Styrax benzoin in Java and

Sumatra are similar to those of A. bambucifoliae in Taiwan. Alatae (BL
1.3-1.7 mm) with secondary rhinaria distributed ANT III 10-22, IV 6-14, V
4-9) emerge through a circular opening in February-June. Hille Ris Lambers
 Atarsaphis 575

(1953b) suggested that the secondary host generations on bamboos were A.

pallida, but this needs experimental confirmation.

Astegopteryx sumatrana Hille Ris Lambers Galls are coral-like, about 10 cm

in diameter when fully developed, with small openings on distal lobes, arising
from the stem via a short thick branching tube (Fig. 124F). They are found
on Styrax spp. (serrulatus var. mollissimus, subdenticulatum) in Sumatra.
Alate emigrants, BL about 1.5-1.6 mm, were collected in February (Hille Ris
Lambers, 1931). Possibly this species belongs in the genus Tuberaphis, in
which case the secondary host would probably be in Loranthaceae.

Astegopteryx unimaculata Noordam Apterae very broad-bodied, pale

yellow or greenish-yellow, with an extensive dark bluish-green dorsal patch,
broadest anteriorly, and a small marginal fringe of white wax; BL 1.0-1.9 mm
(Noordam, 1991). On lower sides of leaves of bamboos (Bambusa, Dendro-
calamus, Gigantochloa) in India, China, Taiwan, Indonesia and Malaysia. In
Taiwan this species seems to be restricted to Dendrocalamus latiflorus (Liao,
1976; as A. insularis). Mounted specimens are difficult to distinguish from
A. bambucifoliae, but in life the broad abdomen and dark green patch are
distinctive. 2n = 12 (Chen and Zhang, 1985b; as A. insularis).

Astegopteryx vandermeermohri (Hille Ris Lambers) Galls are extensive

structures consisting of very long, thin, dichotomously branched tubes attain-
ing 20-35 cm in length, resembling some species of stagshorn fern (Fig. 124H).
On Styrax spp. (serrulatus var. mollissimus, subdenticulatum) in Sumatra.
The emigrant alatae, BL about 2.1 mm, were collected in February-March.
The secondary host is unknown. The generic placement of this species is
uncertain (S. Aoki, pers. comm.). (N.B. Trichoregma vandermeermohri
Takahashi, 1935a, described from bamboos, is a synonym of Astegopteryx
basalis.)

ATARSAPHIS Takahashi Hormaphidinae: Nipponaphidini

Only one species is known, characterized by the subdivision of the dorsal

cuticle into ornamented plates, the lateral outgrowths of the venter and the
extreme reduction of antennae and tarsi in later instars and adults.

Atarsaphis agrifoliae (Ferris) Anholocyclic, feeding throughout the year

on young branches and along petioles and mid-ribs of evergreen oak leaves
(Takahashi, 1958b, as A. quercus). Developmental stages are fully described
by Sorin (1961). Apparently no alate morphs or sexuales are ever produced.
Only found on Quercus phylliraeoides in Japan, where it is presumably native,
but it is also recorded, and was first described, from California, USA, on
Q. agrifolia (Hille Ris Lambers, 1966d).
576 Aulacophoroides

AULACOPHOROIDES Tao Aphidinae: Macrosiphini

A genus for 2-3 species in East Asia on Leguminosae, with a spinulose,

Aulacorthum-like head but also with characters similar to Megoura. (The
generic classification of the oriental legume-feeding Macrosiphini needs to be
clarified).

Aulacophoroides formosanum (Takahashi) Apterae with BL about 2.7 mm,

green with yellowish-brown head, black antennae and siphunculi, yellowish-
brown legs except for black tarsi and a yellow CAUDA. Recorded only from
Millettia reticulata in Taiwan (Takahashi, 1923) and Chekiang, China (Tao,
1963).

AULACORTHUM Mordvilko Aphidinae: Macrosiphini

A genus of about 50 species characterized by the steep-sided inner faces of

the spinulose antennal tubercles. They are mostly monoecious on herbaceous
plants, without any clear pattern of host plant relationships. The nine species
recorded from trees include both polyphagous aphids (circumflexum, mag-
noliae, solani) and some that are highly host-specific. See also B & E, 1984,
p. 239.

Aulacorthum cercidiphylli (Matsumura) Apterae with BL 1.8-2.1mm in

spring, but down to 1.1-1.2 mm in summer, yellow to pale green in life,
feeding singly on leaves of Cercidiphyllum japonicum in Japan (Honshu,
Hokkaido). Presumably monoecious, holocyclic. The alata is described by
Shinji (1930), and the fundatrix by Miyazaki (1971). 2n = 12.

Aulacorthum (Neomyzus) circumflexum (Buckton) Apterae with BL 1.2-

2.6 mm, shining pale yellow to bright green, with a characteristic horseshoe-like
mark on the dorsal abdomen. Anholocyclic and cosmopolitan on numerous
plants, including both monocots and dicots, and also sometimes on gymno-
sperms (Picea, Pinus). See B & E, 1984, p. 239.

Aulacorthum ibotum Essig and Kuwana Apterae with BL 1.8-2.3 mm,

yellow to pale green with dark antennae, yellowish legs except for black tarsi,
black siphunculi and a pale CAUDA (Essig and Kuwana, 1918). On undersides
of leaves of Ligustrum obtusifolium (= ibota) in Japan (Hokkaido, Honshu).
Life cycle has not been investigated. 2n = 14.
 Aulacorthum 577

Aulacorthum linderae (Shinji) Apterae with BL 2.0-2.5 mm, pale green

to white or creamy with brownish coloration around bases of siphunculi;
antennae and legs banded with black, siphunculi pale with black tips, CAUDA
pale. In Japan, on undersides of leaves of Lindera sericea, causing loose leaf
curl and reddish-brown blotches (Miyazaki, 1971). Presumably monoecious,
holocyclic. (The host was given as Aucuba japonica in the original description
(Shinji, 1922b), presumably in error.)

Aulacorthum magnoliae (Essig and Kuwana) Apterae with BL 2.5-3.0 mm,

distinctly coloured with reddish head and prothorax, rest of body yellow-green
to green; femora mostly black, tibiae and antennae usually wholly black,
siphunculi pale with dark tips, CAUDA dark. Polyphagous and mainly
anholocyclic, in Japan, China, Korea and India (B & E, 1984, p. 241); with
a 'relict' holocycle on Sambucus in Japan. Matsuka and Imanishi (1982)
studied its life cycle on Sambucus sieboldiana near Tokyo, where populations
overwinter both as viviparae and, less commonly, as eggs. Clones descended
from fundatrices produced males and viviparous females, but very few
oviparae, so the sexual phase was almost non-existent in that population.
Males are also recorded from India (Raychaudhuri et al., 1980b). 2n = 12.

Aulacorthum muradachi (Shinji) Apterae with BL 2.0-2.3 mm, pale yellow

to green with reddish-brown coloration at bases of the black siphunculi, on
leaves of Lindera spp. (erythrocarpa, praecox, triloba, umbellata) in Honshu,
Japan. Presumably monoecious holocyclic; fundatrices are described from
L. triloba (Takahashi, 1965a; Miyazaki, 1971). 2n = 10.

Aulacorthum rhamni M.R. Ghosh, A.K. Ghosh and Raychaudhuri Apterae

with BL about 1.8-2.0 mm, yellowish-green with darker siphunculi and
CAUDA. On apical leaves of Rhamnus nepalensis in North Bengal, India
(M.R. Ghosh et al., 1971). Biology is unknown.

Aulacorthum solani (Kaltenbach) Apterae with BL 1.8-3.0 mm, colour

variable; rather shiny, whitish-green to yellow, with green or reddish colora-
tion at bases of siphunculi, or uniformly green to brown. Alatae are of much
darker appearance. Cosmopolitan and very polyphagous on both dicots and
monocots, including many trees (B & E, 1984, p. 241). 2n = 10.

Aulacorthum syringae (Matsumura) Apterae with BL about 2mm, pale

yellow to yellowish-green, often with reddish-brown coloration around bases
of siphunculi. Antennae and legs mainly dark, siphunculi black, CAUDA pale
(Takahashi, 1965a). On Syringa spp. (amurensis, emodi, reticulata, vulgaris)
in Japan. Presumably monoecious, holocyclic; fundatrices are described from
S. reticulata (Miyazaki, 1971).
578 Avicennina

AVICENNINA Narzikulov Aphidinae: Macrosiphini

A genus with nominally two species. The type species is described from
Lonicera, and has first tarsal segments with 3 hairs, elongate siphunculi and
alatae without rhinaria on antennal segment IV. The Indian Prunus-feeding
species placed here has 5 hairs on first tarsal segments, rather short siphunculi
and numerous rhinaria on antennal segment IV of alatae, suggesting that it
may not belong in this genus.

Avicennina indica Chakrabarti and Maity Apterae of BL about 4.6-4.9 mm,

pale greyish covered with powdery wax, with brown, clavate siphunculi.
Alatae have black head and thorax and blackish-brown dorsal, abdominal
markings. Fundatrices are greenish with dark dorsal spots. It feeds along the
mid-ribs on upper sides of leaves of Prunus cornuta in late May-June, causing
the leaf laminae to fold upwards and come together above the aphids. In
Uttar Pradesh, India (Chakrabarti and Maity, 1984). Heteroecious holocyclic;
alatae leave Prunus in late June to fly to an unknown secondary host (Medda
et al, 1986).

BAIZONGIA Rondani Pemphiginae: Fordini

A genus for one or two species related to Aploneura and Slavum but with more
normal distribution of rhinaria on the antennae of the alatae, which fold their
wings roof-like in repose. The large characteristic horn-like galls (Fig. 126C)
can be formed on several species of Pistacia.

Baizongia pistaceae (Linnaeus) Producing elongate horn-like galls 15-22 cm

long on Pistacia spp. in the Mediterranean area, and also in northwest India.
Large numbers of alatae, BL 1.8-2.0 mm, emerge in September-November
and fly to roots of Gramineae. The holocycle takes two years (Fig. 2).
Anholocyclic populations occur on grass roots more widely in Europe, Africa
and the Indian subcontinent (B & E, 1984, p. 242). Roberti (1983) and
Wertheim (1954) give accounts of this aphid in Italy and Israel respectively.
Wool (1990) analysed data from a ten-year population study on Pistacia in
Israel. 2n = 24. (A second species, B. yunlanensis, host unknown, is described
from China (Zhang and Zhong, 1985b).)
 Betacallis 579

BETACALLIS Matsumura Drepanosiphinae: Phyllaphidini

Six species are known, all from East and Southeast Asia. All viviparae are
alate, active insects (behaving like Euceraphis), and have a characteristic
transverse dark brown to black bar across the ventral side of the head between
the eyes. Most of the species are recorded from Betulaceae, which are also the
hosts of several related genera. The Indian species are reviewed by Chakrabarti
(1988) and A.K. Ghosh and Quednau (1990), and the Japanese species by
Higuchi (1972).

Betacallis alnicolens Matsumura Alatae are pale green with dark tibiae,
large (about 3.5mm), very active, on leaves of Alnus spp. (hirsuta, japonica,
matsumurae) in Japan, Korea and China, and also collected from Corylus
mandshurica in Korea (BMNH colln). 2n = 22.

Betacallis luminiferus Zhang Alatae of BL about 2.6mm, on leaves of

Betula luminifera in Hangzhou, China. Very similar to, and possibly synony-
mous with, B. sikkimensis, but with more secondary rhinaria.

Betacallis odaiensis Takahashi Alatae are pale lemon yellow to apricot, with
black basal, sensoriated, part of third antennal segment, black tibiae, two
black bars on the dorsal abdomen, and black siphunculi; BL 2.4-3.1 mm. On
leaves of Betula spp. (ermannii, maximowicziana, platyphylla var. japonica)
in Japan. (Indian records of odaiensis refer to other species.) 2n = 22.

Betacallis prunicola R.C. Basu, A.K. Ghosh and Raychaudhuri Alatae light
yellowish-green with dark tibiae, dark transverse bars on abdominal tergites
4 and 5 and dark siphunculi; BL 1.9-2.2 mm (A.K. Ghosh and Quednau,
1990). Described from adults and immatures on Prunus cerasus in Meghalaya,
India (R.C. Basu et al., 1975), but this is unlikely to be the true host. This
species is also recorded from Arunachal Pradesh, Manipur and West Bengal,
and from China and the Philippines. It can be distinguished from other
Betacallis species by the long ultimate rostral segment (R IV+V 1.5-1.7 x
HT II).

Betacallis querciphaga R.C. Basu, M.R. Ghosh and Raychaudhuri Alatae

light grenish-yellow with dark markings like B. odaiensis except for the dorsal
longitudinal head stripe and differences in wing vein pigmentation; BL 2.4-
3.2 mm. In India (Manipur, Meghalaya, Sikkim, West Bengal), the Philippines
(Chakrabarti, 1988) and Thailand (BMNH colln). It was described from
Quercus sp., and alatae and immatures are also recorded from Castano-
spermum sp., so Fagaceae may act as reserve hosts; nevertheless, specimens
from Betula alnoides in Thailand and from B. utilis in West Bengal (BMNH
580 Betulaphis

colln) indicate that the normal host association is probably with Betula.
Parasitized by Trioxys (Betuloxys) assamensis in India (D. Raychaudhuri,
1990).

Betacallis sikkimensis R.C. Basu, M.R. Ghosh and Raychaudhuri BL of

alata 2.4-3.2 mm, appearance in life unrecorded but presumably pale, with
third antennal segment and tibiae darkened only at apices (cf. other species).
The siphunculi are black on distal half only and there is often a consolidated
or broken dark central patch on abdominal tergites 4 and 5. Described from
Quercus sp. - which is probably only a reserve host - in Sikkim, India (R.C.
Basu et al., 1974); since collected many times in Uttar Pradesh from Betula
alnoides and Betula sp. (Chakrabarti, 1988; and BMNH colln). Chakrabarti
(1988) described the alate male and apterous ovipara from Betula. 2n = 20
(Khuda-Bukhsh and Pal, 1983b).

BETULAPHIS Glendenning Drepanosiphinae: Phyllaphidini

A genus of about eight or nine species rather evenly distributed through

the holarctic on Betula. They are small, rather flat oval aphids with short
6-segmented antennae, the terminal process being about as long as the base
of segment VI. Viviparae have a short, conical CAUDA and a bilobed anal
plate. Oviparae have dark dorsal markings and the posterior abdomen is
extended as an 'egg-laying organ'. Males are apterous with extensive dark
dorsal markings. Fundatrices are unusual in being either apterous or alate (at
least in B. quadrituberculata). Stroyan (1977) gives a detailed generic diagnosis
and accounts are available for North America (Richards, 1961), Canada
(Richards, 1969b), Fennoscandia and Denmark (Heie, 1982) and India
(Chakrabarti, 1988; A.K. Ghosh and Quednau, 1990). There has been much
confusion about the number of species in the brevipilosa/quadrituberculata
group, due to seasonal variation in hair lengths. Here, B. helvetica HRL is
regarded as a synonym of B. brevipilosa, although a study of haemolymph
proteins by Karwanska (1987) suggests that it may be a distinct species.
Parasitized by Aphidius aquilus, which also attacks other related aphids
feeding on Betula (Stary, 1973).

Betulaphis brevipilosa Börner (= helvetica Hille Ris Lambers) Apterae are

pale to grass-green aphids, BL 1.5-2.0 mm, with the tips of the antennae and
the tarsi dark. Alatae almost always have a dark dorsal abdominal patch. This
species is frequently confused in the literature with B. quadrituberculata. It
seems to feed exclusively on Betula pendula, usually on the upper surfaces of
the leaves, in northern and Central Europe (but not the UK), and introduced
into North America and New Zealand. Accounts are given by Cottier (1953,
as B. quadrituberculata) and Heie (1982). Ecology of introduced populations
 Betulaphis 581

in northern California was studied by Hajek and Dahlsten (1988). Betulaphis

viridis Richards, 1969b seems to be this species. 2n = 20.

Betulaphis hissarica Narzikulov Appearance in life is unknown. BL of

aptera about 1.7-1.9 mm. Chakrabati (1988) redescribed both apterous and
alate viviparae, and also apterous oviparae and males. Recorded from various
Betula species (alnoides, hissarica, turkestanica) in Tadzhikistan and India
(Himachal Pradesh, Uttar Pradesh).

Betulaphis japonica Takahashi Appearance in life and biology unrecorded,

and alatae are unknown. BL of aptera 1.1-1.6 mm. Found on Betula platy -
phylla var. japonica and B. maximowicziana in Japan, on B. platyphylla in
Korea, and also recorded from B. fruticosa in Mongolia. For further details
see Higuchi (1972).

Betulaphis longicornis Quednau and Chakrabarti Apterae yellow: BL 2.3-

2.9mm. The alatae lack a dorsal abdominal dark patch, and have numerous
(26-35) closely-spaced elongate transverse secondary rhinaria on antennal
segment III. Apterous and alate viviparae are described by Quednau and
Chakrabarti (1980), and apterous oviparae and males by Chakrabarti (1988).
Recorded from Betula alnoides and B. utilis in India (Assam, Bhutan,
Himachal Pradesh, Uttar Pradesh, West Bengal).

Betulaphis pelei Hille Ris Lambers Apterae are yellow with dark tarsi
and tips of antennae, BL 1.4-1.7 mm. Alatae sometimes have a dark dorsal
abdominal patch. On Betula nana with a boreo-alpine distribution in northern
Europe, Iceland and Greenland (Heie, 1982). Aphids occurring on Betula
glandulosa, described from Baffin Island as Betulaphis arctosetis Richards,
1961, are probably also this species. 2n = 20.

Betulaphis quadrituberculata Kaltenbach Apterae have BL 1.5-2.0 mm and

are pale yellowish-green or pale yellow to almost white, sometimes mottled
with darker pigment, normally feeding on the undersides of leaves. Alatae
sometimes have a dark dorsal abdominal patch. Recorded from various Betula
species, especially B. pubescens and also occasionally found on Alnus incana.
Widely distributed in Europe (including Iceland) and across Asia to Mongolia,
and introduced into North America. There is much variation in length of
the spinal and pleural hairs on the anterior abdominal tergites which causes
short-haired specimens of this species, found particularly in the summer
months, to be confused easily with B. pelei and B. brevipilosa. Betulaphis
aureus Richards, 1961, described from Betula sp. in Canada, seems to be this
short-haired form of B. quadrituberculata. Varty (1964) has studied its
biology in New Brunswick, Canada. See also Heie (1982), Stroyan (1977).
2n = 20.
582 Boernerina

BOERNERINA Bramstedt Drepanosiphinae: Phyllaphidini

A genus of four to five species associated with Alnus (including Duschekia).

The apterae are dorsoventrally flattened, rather elongate oval, and apply
themselves closely to the leaf surface. They have paired anteriorly-directed
rectangular projections on the head, lateral abdominal hairs arising from
asymmetric tubercles that give the abdominal margin a stepped appearance
and typically 5-segmented antennae. The alatae are more normal looking
Phyllaphidini. One species (occidentalis) differs in that the apterae often have
6-segmented antennae, the frontal projections are only weakly developed and
the abdominal margin is more indented than stepped. Hille Ris Lambers and
Hottes (1962) revised the genus.

Boernerina alni Takahashi Apterae are pale yellowish-white, dorsoventrally

flattened, elongate oval, BL 1.7-2.5 mm. Alatae are whitish and waxy. Sorin
(1970) described the apterous male and ovipara, collected in mid-August
in Japan, on Alnus spp. Pashchenko (1988b) has described a subspecies
(insularia) in the former Soviet Far East.

Boernerina depressa Bramstedt Apterae are whitish, flattened and elongate

oval, BL 1.5-2.5 mm, on undersides of leaves of Alnus viridis in the Alps and
Carpathians. Oviparae and very small apterous males are produced in July-
August (Hille Ris Lambers and Hottes, 1962).

Boernerina (Boernerinella) occidentalis Hille Ris Lambers and Hottes

Apterae are pale, flattened, oval, BL 1.7-2.4 mm, with broad sclerotic bars
on abdominal tergites, the anterior ones being sometimes brownish pigmented.
Alatae are variably pigmented, darker specimens have blackish-brown sclero-
tic bars on abdominal tergites. On Alnus sinuata in Alaska, and also collected
in Oregon (BMNH colln; leg. DHRL). Oviparae and small, slender apterous
males are produced in July in Alaska (Hille Ris Lambers and Hottes, 1962).

Boernerina variabilis Richards Apterae are pale to bright yellow with black
on distal parts of antennae and legs, flattened, elongate oval, BL 1.9-2.6 mm.
Alatae vary in pigmentation; when well pigmented they have broad dark bars
dorsally and ventrally on the abdomen. On undersides of leaves of Alnus spp.
(sinuata, rugosa, alnobetula) in east and west Canada (British Columbia,
Northwest Territory, Ontario, Quebec, Labrador). Oviparae and small,
apterous males occur in August in Northwest Territory (BMNH colln, leg. R.
O'Doherty), but oviparae were collected in October in northern Ontario
(BMNH, leg. J. Sÿpkens). 2n = 16*.
 Brachycaudus 583

BRACHYCAUDUS van der Goot Aphidinae: Macrosiphini

This genus is characterized by the large round spiracular apertures, short

often semicircular CAUDA and siphunculi with a subapical annular incision.
Primitively heteroecious with Prunus as primary hosts and about 14 species
still host-alternate. The phylogeny of Brachycaudus in relation to its host
plants is discussed by Shaposhnikov (1956). For an introduction to the
genus and more detailed accounts of seven of the Prunus-feeding species see
B & E, 1984, p. 243. Heie (1992) reviewed the species in Fennoscandia and
Denmark.

Brachycaudus almatinus Nevsky Apterae according to the original descrip-

tion are pale brown with black lateral spots, black siphunculi, femora, tibial
apices and tarsi, and dusky antennae. BL about 1.8 mm. The CAUDA is very
short and broad. Alata have a pale brown abdomen, presumably without a
dark dorsal patch. Found in Kazakhstan (Alma-Ata) on plum and pear, rolling
leaves longitudinally (Nevsky, 1951). Biology unknown.

Brachycaudus (Thuleaphis) amygdalinus (Schouteden) Host-alternating

between Prunus dulcis or P. persica and Polygonum spp. Apterae are green
with variably developed dark dorsal markings, squat-bodied with rather short
pale legs and antennae; BL 1.6-2.1 mm. Alatae have a dark central abdominal
patch. Colonies in spring cause rolling of young almond or peach leaves and
stunting of new growth. The leaves are rolled somewhat obliquely with respect
to the mid-rib, rather than perpendicularly to the mid-rib as in B. helichrysi.
In Europe, the Middle East, Pakistan and South Africa. Darwish (1983)
described the morphology of the spring generations. See also B & E, 1984,
p. 243. 2n = 12.

Brachycaudus cardui (Linnaeus) Host-alternating between Prunus domestica

or P. spinosa (or sometimes P. avium or P. armeniaca) and various Com-
positae and Boraginaceae. Apterae of BL 1.9-2.3 mm, in rolled leaves of plum
in spring, are usually green with the dorsal cuticle sclerotic and often brownish
but not shiny black like the summer generations on secondary hosts. Alatae
have extensive dark dorsal pigmentation. In Europe, North Africa, southwest
and Central Asia, India, China and North America. According to Börner
(1952-3), the aphids which use P. spinosa as primary host and migrate to basal
parts of stems of Compositae are a distinct species, B. lateralis (Walker), but
further work seems necessary to confirm this. See also B & E, 1984, p. 243.
2n = 10.

Brachycaudus divaricatae Shaposhnikov Host-alternating between Prunus

divaricata (sometimes P, domestica or P. spinosa) and Caryophyllaceae
584 Brachycaudus

(Melandrium album) in Eastern Europe and the Middle East (northern

Caucasus, Transcaucasia, Crimea, Turkmenia, Turkey, Iran). Apterae are
black, BL 2.0-2.2 mm, rolling leaves of Prunus in spring; the migration to
Caryophyllaceae is apparently facultative (Shaposhnikov, 1962).

Brachycaudus divaricatellus Shaposhnikov Apterae of BL about 2.0mm,

shiny yellowish-rose, elongate oval, on undersides of leaves of Prunus
divaricata, causing slight curling of tips and discoloration. In southwest
Turkmenia (Shaposhnikov, 1964a). Life cycle is unknown. Shaposhnikov
(1964b) described the ovipara.

Brachycaudus helichrysi (Kaltenbach) Host-alternating between Prunus

spp. (domestica, insititia, spinosa) and numerous secondary hosts, especially
herbs in Compositae and Boraginaceae. Of worldwide distribution; anholo-
cyclic populations occur in warmer regions and are sometimes found on new
growth of various trees. Apterae in spring populations on Prunus are green,
brownish or yellowish, shiny and slightly waxy, causing leaves to roll up tightly
perpendicular to the mid-rib. BL 1.4-2.0 mm on Prunus, down to 0.9 mm on
secondary hosts. Darwish (1984) described the morphology of spring genera-
tions; for other information see B & E, 1984, p. 244. 2n = 12.

Brachycaudus persicae group Apterae shiny dark brown or black, BL 1.5-

2.2mm, with black siphunculi. Brachycaudus persicae (Passerini) is most
commonly found in large spring colonies on young stems of Prunus persica
or P. armeniaca, often persisting into the summer on root suckers. Host alter-
nation occurs to aerial parts of Scrophulariaceae (Euphrasia, Melampyrum,
Rhinanthus) in Europe (Burger, 1975), but anholocyclic overwintering is also
common on Prunus roots. Aphids commonly regarded as this species are also
widely distributed on peach outside Europe, in the Middle East, southern
Africa, Australia, New Zealand, North and South America.
Similar aphids also occur commonly in Europe on P. domestica and
P. spinosa, and occasionally on Pyrus communis and Chaenomeles japonica.
These all form a complex of which the host relationships and life cycles
need further clarification. Müller and Steiner (1988) applied the name
B. semisubterraneus Börner to anholocyclic populations on plum in Germany,
and suggested that the form distributed worldwide on peach was also
an anholocyclic species, to which they applied the name B. persicaecola
(Boisduval). However, aphids of the B. persicae group are recorded from
Scrophulariaceae in both Australia and Chile (BMNH colln), indicating that
some host alternation occurs in southern continents, and Burger (1975)
showed that not all populations on plum are anholocyclic, so the application
of the names semisubterraneus and persicaecola is still unclear. 2n = 10.

Brachycaudus (Mordvilkomemor) pilosus (Mordvilko ex Nevsky) (= cerasicola

Mordvilko ex Nevsky) ( = phlomicola Nevsky) Apterae are brown with an
extensive black dorsal shield; BL 1.8-1.9 mm. Immatures are dirty brown.
Spring colonies roll and twist the leaves of Prunus spp. and turn them red.
 Brachyunguis 585

Primary hosts include Prunus bucharica, P. erythrocarpa, P. microcarpa,

P. tianschanica, P. ulmifolia. Damaged leaves of P. ulmifolia develop gall-
like thickenings and red spots (Narzikulov, 1965b). There is probably a
facultative host alternation to Labiatae (Shaposhnikov, 1964b; as phlomicola).
In mountainous regions of Central Asia: Kazakhstan, Kirgizia, Tadzhikistan,
Turkmenia, and Pakistan (BMNH colln).

Brachycaudus (Appelia) prunicola (Kaltenbach) Apterae of BL 1.4-2.4 mm,

shiny black, dark green or deep yellow-green with black dorsal markings;
immatures grey-green. In spring on new growth of Prunus spp. (usually
P. spinosus, sometimes P. domestica or P. insititia), causing severe leaf curl
and discoloration. Host-alternating between Prunus and Tragopogon, but
part of the population - possibly a subspecies - may live without alternation
on P. spinosus. Not yet recorded outside Europe. 2n = 12.

Brachycaudus (Appelia) schwartzi (Börner) Apterae of BL 1.4-2.1mm,

shiny yellow-brown to dark brown with extensive dorsal black sclerotization;
immatures are yellow-brown. Living without host alternation on Prunus
persica; spring colonies cause severe curling and distortion of peach leaves.
Prunus serotina is the only other recorded host. In Europe, Iran, India, South
America and California. Records of B. tragopogonis from peach in the
literature are this aphid. Darwish (1983) gives descriptions of all life cycle
stages. (See also B & E, 1984, p. 246). 2n = 12.

BRACHYSIPHONIELLA Takahashi Aphidinae: ?Aphidini

One species. Brachysiphoniella montana (van der Goot), with short siphunculi
and a long CAUDA, mostly known from wax-covered colonies on aquatic
Gramineae (Takahashi, 1923) in East and Southeast Asia, and Australia. In
Korea there appears to be host alternation, with Pyrus montana as the primary
host, spring colonies causing leaf curl (Paik, 1972); as Semiaphis montana).
However, this needs confirmation, as it is possible that Paik's account con-
fuses B. montana with Melanaphis siphonella.

BRACHYUNGUIS Das Aphidinae: Aphidini

Related to Aphis, but with antennal processus terminalis clearly shorter than
base of last segment. The siphunculi are pale and much shorter than the
CAUDA, and the body is usually covered with wax meal or dust. There are
about 25 nominal species, mostly associated with xerophytes in Central and
586 Brachyunguis

southwest Asia (Nevsky, 1928b; as Xerophilaphis; Bodenheimer and Swirski,

1957). The genus is poorly known; many of the nominal species are very similar
to one another and their separate identity needs experimental confirmation.

Brachyunguis calligoni (Nevsky) Apterae are light green covered with wax
meal, BL 1.3-1.7 mm, with very small inconspicuous siphunculi, living on
young shoots of Calligonum spp. (eriopodium, setosum, ?comosum) in
Uzbekistan (Nevsky, 1928b). Biology is unknown.

Brachyunguis calotropicus Menon and Pawar Apterae pale to muddy green,

with sparse wax powder, BL 1.2-1.5 mm. In dense aggregations on upper or
undersides of younger leaves of Calotropis procera, sometimes attended by
ants. Life cycle is unknown. First recorded from the Delhi region of India
without a proper description (Menon and Pawar, 1958). Brachyunguis ushari
Müller and El Tigani, 1986 seems to be a synonym, and the present account
is based on that assumption. In India, southwest Asia (Saudi Arabia) and
Africa (Algeria, Egypt, Sudan). 2n = 8 (Kurl, 1978).

Brachyunguis harmalae Das Apterae are mid- to deep green anteriorly,

more yellowish on posterior abdomen, thickly dusted with wax powder,
and with two longitudinal rows of submarginal dark green spots; BL 1.2-
1.8mm. Often ant-attended. Monoecious holocyclic on Peganum harmalae in
Pakistan, where Das (1918) studied its life cycle in detail. Both apterous and
alate males occur. The populations that occur throughout the southwest Asia,
in Africa (Eritrea, Kenya, Sudan, Senegal) and on the Iberian peninsula,
colonizing various, mostly xerophytic plants including Calligonum and
Calotropus, are presumably anholocyclic. Eastop and Raccah (1988) list 12
hosts in seven botanical families in the Arava valley, Israel. An outbreak is
also recorded on Citrus (Swirski, 1963). Brachyunguis plotnikovi (q.v.) in
Central Asia is probably this species.

Brachyunguis plotnikovi Nevsky Monoecious holocyclic, with alate males,

on Calligonum spp. (aphyllum, arborescens, comosum, eriopodium, setosum)
in Central Asia (Kazakhstan, Turkmenia, Uzbekistan). Viviparae are recorded
from plants in several families and damaging populations can build up on
cotton (Nevsky, 1928b; as Xerophilaphis). The description of this species does
not distinguish it from B. harmalae. Only alate males were recorded, and no
ant attendance was observed.

Brachyunguis tamaricis (Lichtenstein), Plate 13a, b Apterae according to

the original description are velvety grey-green, with no mention of wax dust
(but development of wax may depend on age and microclimate); BL is
0.9-1.7 mm. Colonies on twigs are inconspicuous, resembling small leaves or
leaf-scales of the host plant. They may be attended by ants (Monomorium).
On Tamarix spp. in southern and Central Europe, North Africa, southwest
and Central Asia east to Pakistan. Life cycle is unknown; Ivanoskaya (1956)
described the fundatrix on T. meyersi in Turkmenia (as tamaricophila), and
 Brysocryptoides 587

specimens from Pakistan on T, kotschyi in the BMNH collection also seem

to be fundatrices. No sexuales have been recorded.
There is much confusion in the literature on the nomenclature of the
Tamarix-feeding species of Brachyunguis. Brachyunguis tamaricophila of
Ivanoskaya (1956; nec. Nevsky, 1928b) seems to be tamaricis, and B. tamari-
civorum Narzikulov) (= tamariciarum Ivanoskaya) is not clearly distinct.
Brachyunguis tamaricifoliae (Hall, 1926) is also a synonym of tamaricis.
Records of tamaricis from Calotropis procera probably refer to either B.
harmalae or B. calotropicus.

Brachyunguis tamaricophila (Nevsky) Apterae according to the original

description are green, densely covered with mealy wax secretion; BL 0.7-
1.7mm. In large, ant-attended colonies on young shoots, leaves and flowers
of Tamarix spp. Monoecious holocyclic in Uzbekistan (Tashkent), oviparae
being found, but males are still undescribed (Nevsky, 1928b). Ivanoskaya
(1956) described the fundatrix on T. karakalensis in Turkmenia (as tamari-
cifoliae). Also recorded from Israel, Libya and Pakistan (BMNH colln).
Brachyunguis tamaricifoliae of Ivanoskaya (1956; nec. Hall, 1926) seems to
be this species.

BREVICORYNELLA Nevsky Aphidinae: Aphidini

A genus with a single species resembling Brachyunguis but with protuberant

eyes lacking ocular tubercles.

Brevicorynella quadrimaculata Nevsky Apterae brown or dark brown,

covered with a fine pruinose secretion, antennae pale yellow with articulations
and all of last segment black, abdomen with four blackish-brown dorsal spots;
BL 1.1-1.6 mm. The aphids cluster on the ends of shoots of Tamarix spp. and
are attended by ants (Crematogaster). Recorded only from Central Asia
(Kazakhstan, Tadzhikistan) (Nevsky, 1928b).

BYRSOCRYPTOIDES Dzhibladze Pemphiginae: Eriosomatini

Three species, two described from galls on the primary host (Zelkova) and
one only known from Cyperaceae. Alate spring migrants, emerging from
pseudogalls on Zelkova, have an unbranched media, only one oblique vein in
the hind wing and inconspicuous ring-like siphunculi. Resembling Kalten-
bachiella in many characters, but in Byrsocryptoides the secondary rhinaria
are less annular and the processus terminalis is relatively longer. Dzhibladze
588 Calaphis

(1965b) provides keys to fundatrices, apterous fundatrigeniae and spring

migrants of the species on Zelkova.

Byrsocryptoides zelkovae Dzhibladze The galls (pseudogalls) are formed by

rolling, distortion and blistering of a terminal or subterminal leaf, starting
at the base, on Zelkova acuminata and Z. carpinifolia in Russia (Georgia;
Dzhibladze, 1960). Alatae leaving the galls in June are small (BL about 1.2-
1.3mm), rather long-bodied, with dark brown head, meso- and metathorax
and pale yellow to green prothorax and abdomen. They probably fly to aerial
parts of Cyperaceae - apterae of elongate form with a long last rostral seg-
ment collected on Carex riparia in Germany (BMNH colln, leg. F.P. Müller)
are provisionally assigned to this species - but the life cycle needs experimental
confirmation.

Byrsocryptoides zelkovaecola Dzhibladze Galls (Fig. 125A, p. 655) are

similar to those of B. zelkovae, and occur on Zelkova carpinifolia in Georgia.
Alatae of BL 1.1-1.4 mm emerge in June-July. Life cycle is unknown
(Dzhibladze, 1965b).

CALAPHIS Walsh Drepanosiphinae: Phyllaphidini

Delicately built aphids with rather long thin legs, generally pale-coloured but
sometimes with distinctive markings and/or dark wing veins. Other characters
of the genus are the well-developed antennal tubercles and the absence or
indistinctness of the radial sector in the forewing. Some species have all
viviparae alate, others have both apterous and alate viviparae. As treated here,
Calaphis is holarctic and includes Kallistaphis and Cepegillettea. There are
about 15 species, 11 of which are in North America, two are European and
one is East Asian. All except one (myricae) live on Betulaceae. Parasitized by
Aphidius aquilus, which also attacks other related Betula-feeding aphids
(Stary, 1973).

Calaphis alni Baker Apterae and alatae of BL about 2.3-2.4 mm, yellowish
with distinctive black markings; alatae usually have a large black dorsal
abdominal patch and apterae have two bands across the thorax and one across
the posterior abdomen. Feeding mainly on the stems and foliage of tender
growing terminals of Alnus rugosa (Pepper, 1950), in eastern USA and
Canada (New Brunswick). The sexuales are undescribed.

Calaphis alnosa Pepper Apterae of BL 1.3-1.6 mm, creamy-white with a

reddish-brown line along each side of the body; antennae and legs dark. They
position themselves close to the larger veins on the undersides of leaves of the
host (Alnus rugosa) and are very inconspicuous. Alatae are very pale green
 Calaphis 589

with dark antennae and legs. Sexuales occur in October; males are alate
(Pepper, 1950).

Calaphis arctica Hille Ris Lambers Apterae of BL about 1.9-2.1mm,

green, with dusky dorsal hairs and spots (O. Heikinheimo, pers. comm.).
Originally described from Betula nana in Greenland, and since recorded
from Finland (BMNH colln; leg. Heikinheimo) and alpine Germany (BMNH
colln, including undescribed alate viviparae). Apterous males and oviparae
were collected in late October in Germany (BMNH colln, leg. DHRL).
2n = 18*.

Calaphis betulaecolens (Fitch) Viviparae all alate, bright lemon yellow

without dorsal markings, with blackish antennae and tibiae and dark wing
veins; BL 3.0-3.5 mm. Usually on leaves of Betula lutea and B. papyrifera,
occasionally on other Betula spp. Widely distributed in North America.
(Japanese records of this species apply to C. similis, q.v.). Quednau (1971)
fully redescribed this species. Apterous oviparae and alate males occur in
September-October. 2n = 20.

Calaphis (Cepegillettea) betulaefoliae (Granovsky) Apterae are yellow,

greenish-yellow or green, without dorsal markings, antennae and legs mainly
pale to dusky with black apex to antennal segment 3, femoro-tibial joints and
tarsi, and black tips to the siphunculi; BL 2.9-3.5 mm. Alatae similar, with
yellow-orange head and thorax and dark wing veins. Occurring singly or in
small groups on undersides of leaves or young growth of Betula spp. (glan-
dulosa, occidentalis, papyrifera, pumila); recorded locally in USA (Illinois,
Wisconsin, Colorado), common and widespread in Canada. Prefers shady
positions; very active, dropping when disturbed. Apterous oviparae and alate
males occur in September-October (Granovsky, 1928b).

Calaphis betulella Walsh, Plate 9e Viviparae all alate, yellowish with orange
and black markings, black stripes along the femora and wholly black tibiae
and tarsi; BL 2.0-2.5 mm. Feeds only on Betula nigra, on undersides of leaves.
Apterous oviparae and alate males occur in September-October. In eastern
USA. 2n = 18.

Calaphis betulicola (Kaltenbach) Apterae pale yellowish to green, with

black apices to antennal segments, femoro-tibial joints, tibial apices and
tarsi and dark tips to siphunculi; BL 2.0-2.3 mm. Alatae have darker wing
veins than the rather similar species C. flava. On undersides of leaves of
Betula spp. (davurica, pendula, platyphylla, pubescens, ermannii, mand-
shurica), usually on seedlings and small trees less than l m high (Stroyan,
1977). Holarctic in distribution, from Western Europe across the former
USSR to Japan, and also in USA (California; Quednau, 1971) and Canada
(New Brunswick, Quebec; BMNH colln). Apterous oviparae and alate
males occur in September-November. (Pre-1957 records include C. flava.)
2n = 18.
590 Calaphis

Calaphis coloradensis Granovsky Apterae pale yellow to yellow-green,

tinged with green, sometimes with a very light brownish cast, with dark apices
to antennal segments, femoro-tibial joints, apices of tibiae and tarsi; BL
1.5-1.7 mm. Alatae have both cubitus veins of forewing heavily bordered.
Feeds only on Betula occidentalis, on lower sides of leaves and terminals of
young growth. In western USA and Canada (Alberta, Colorado, Idaho,
Montana, Washington, Utah). Apterous males and oviparae occur in
September-October (Granovsky, 1939; Palmer, 1952). 2n = 18.

Calaphis flava (Mordvilko) (= basalis Stroyan, 1957b; = granovskyi Palmer,

1952) Apterae are pale green or yellowish, with dark apices to antennal
segments, femoro-tibial joints, tibial apices and tarsi; BL 1.9-2.7 mm. Alatae
have wing veins dark, but less dark than in betulicola and the siphunculi are
pale. Betula pubescens seems to be the preferred host in Europe (Heie, 1982),
but the young growth of many other Betula species may be colonized and it
has also been collected from Alnus firma and A. serrulata (BMNH colln).
Widespread in Europe (where until 1957 it was confused with the less common
betulicola), eastward to Siberia and Central Asia, and introduced into South
Africa, Australia, New Zealand and North America (where it was described
as granovskyi - Palmer, 1952). Alate males and apterous oviparae occur in
September-November and may be produced on numerous Betula spp. (BMNH
colln). 2n = 18.

Calaphis leonardi Quednau Viviparae all alate, pale yellowish-green and

orange with brown-black antennae, tibiae and tarsi and dark wing veins; BL
about 3.0-3.3 mm. Feeds only on Betula populifolia, in northeastern USA and
eastern Canada (Quednau, 1971). Oviparae have dark dorsal markings and
occur in September-October; males are not yet recorded. 2n = 20*.

Calaphis manitobensis Richards Apterae are bright yellow, with antennae

and legs almost entirely pale; BL about 2 mm. Alatae and sexuales are
undescribed. Known only from the original description, from Betula pumila
in Manitoba, Canada (Richards, 1968e).

Calaphis (Cepegillettea) myricae (Patch) Apterae and alatae are both green
with dusky or dark distal antennal segments, tibiae and tarsi and siphunculi
either wholly or apically black; BL about 2.5 mm. On leaves of Myricacae
(Myrica asplenifoliae, M. gale, Comptonia peregrina) in northeastern USA
and eastern Canada. Apterous males and oviparae occur in September
(BMNH colln.; leg. E. MacGillivray).

Calaphis neobetulella Quednau All viviparae alate, greenish yellow with

black dorsal markings on head, thorax and abdominal tergites 4-5, and
antennal flagellum, legs and siphunculi also mainly or entirely black; BL
2.0-2.5 mm. On undersides of leaves of Betula nigra in northeastern USA
(North Carolina, Virginia, New Jersey, Washington DC). Alate males and
apterous oviparae occur in September (Quednau, 1971).
 Callipterinella 591

Calaphis similis Quednau All viviparae alate, yellow with reddish head
and two red spots on abdomen, with antennal flagellum, tibiae and tarsi
mainly black, siphunculi pale, wing veins bordered with black; BL 3.0-
3.5 mm. On undersides of leaves of Betula spp. (costata, davurica, mand-
shurica, platyphylla var. japonica) in China, Korea, Siberia and Japan
(Pashchenko, 1988b; Quednau, 1979). Oviparae and males were collected in
October (BMNH colln, and Paik, 1972; as Kallistaphis sp.). Higuchi's (1972)
account of C. betulaecolens applies to this species,

Calaphis viridipallida Palmer Apterae and alatae pale greenish-yellow with

apices of antennal segments, femoro-tibial joints, tibial apices and tarsi dark;
BL 1.6-2.2 mm. On undersides of leaves of Betula spp. (glandulosa, lutea,
papyrifera), widely distributed in North America (California, Colorado,
Minnesota, North Carolina, Manitoba, Ontario, Quebec, Labrador). Oviparae
and alate males occur in September-October. 2n = 18.

CALLAPHIS Walker See PANAPHIS Kirkaldy

CALLIPTERINELLA van der Goot

Drepanosiphinae: Phyllaphidini

A genus of three species associated with Betula. Apterous viviparae occur

commonly and have a brown vertex to the head and usually other dark dorsal
markings. The siphunculi are dark and have rows of minute spinules. Stroyan
(1977) and Heie (1982) both provide accounts of this genus.

Callipterinella calliptera (Hartig) (= annulatus Koch) Apterae yellowish-

green to green, usually with dark transverse bands on all tergites; BL 1.5-
2.5mm. Alatae have dorsal markings less well developed. In ant-attended
groups on undersides of leaves of Betula spp., often inside leaves sewn up by
lepidopterous larvae; B. pendula, less commonly B. pubescens or (in Japan)
B. platyphylla. It occurs throughout Europe and across northern Asia to
Siberia, Korea and Japan; introduced and widespread on planted B. pendula
in North America. Oviparae and alate males occur in September-October.
Hajek and Dahlsten (1988) studied the population dynamics of this species in
California. 2n = 20.

Callipterinella minutissima (Stroyan) Apterae are very small, stout, oval-

bodied, green to yellowish-green with a dark transverse bar on ABD TERG
8 only; BL 0.9-1.4 mm. On Betula spp. (pendula, pubescens) in Europe, and
592 Capitophorus

introduced into western North America, where it is also recorded from

B. papyrifera (BMNH colln). The size and shape of the apterae enable them
to feed inside bud scales and developing leaves when the buds burst in spring,
and in the female catkins in summer (Hajek, 1985). Oviparae (which are larger
and darker than apterous viviparae) and alate males occur on the leaves in
September-November.

Cattipterinella tuberculata (von Heyden), Plate 9c, d Apterae are yellowish

with a brown head and dark dorsal abdominal markings comprising especially
a dark quadratic patch on ABD TERG 4-6; BL 1.7-2.2 mm. Alatae have
irregular small sclerites on posterior abdominal tergites, but no transverse
bands. Apterous males and oviparae are found in September. Apparently
specific to Betula pendula and until recently not recorded outside of Europe.
However, Pashchenko (1988b) records apterous Callipterinella with a dorsal
quadrate patch similar to that of C. tuberculata on various Betula spp. in
Siberia, as well as apterae without dorsal abdominal markings on the same
hosts, indicating that the specific distinction between callipterus and tuber-
culata, which are closely related, may not be applicable in that part of the
world. Also recorded from China (Zhang et al., 1986b). 2n = 20.

CAPITOPHORUS van der Goot Aphidinae: Macrosiphini

About 30 species of pale slender aphids with elongate appendages, and

apterae with at least some of dorsal hairs capitate. Alatae have a dark
dorsal abdominal quadrate patch and numerous rhinaria on antennal seg-
ments III, IV and usually V. Six species are known to have host alternation,
with primary hosts all in Elaeagnaceae and one or more live without host
alternation on Elaeagnus. Secondary hosts are herbaceous Compositae
and Polygonaceae. The genus was onced used in a much broader sense to
include most Macrosiphini with capitate hairs. Taxonomie accounts are
available for North America (Corpuz-Raros and Cook, 1974), Western
Europe (Hille Ris Lambers, 1953a), Central Europe (Heinze, 1961a), India
(Raychaudhuri et al., 1980b) and Japan (Miyazaki, 1971). (See also B & E,
1984, p. 249.)

Capitophorus archangelskii Nevsky Apterae are pale green to yellow-green

with green markings; BL 1.2-1.8 mm, on undersides of leaves of Elaeagnus
spp. (angustifolia, occidentalis) and sometimes persisting on ends of shoots
throughout the year (Nevsky, 1928a). Alatae have a dark green central quadrate
patch on the dorsal abdomen. In Caucasus, Kazakhstan, Afghanistan, Iran
and India (West Bengal, Uttar Pradesh; BMNH colln). Secondary host popu-
lations are unknown; possibly it is monoecious on Elaeagnus. Sexuales occur
in October-November; males are alate (Nevsky, 1928a).
 Capitophorus 593

Capitophorus elaeagni (del Guercio) Apterae in spring populations are pale

green, BL 1.9-2.5 mm, on undersides of leaves of Elaeagnus spp. and some-
times on Hippophae. Fundatrices have brighter green dorsal spots in longitu-
dinal rows and feed on upper sides of young leaves. Alatae, produced in the
second and third generations, have black head and thorax, black antennae and
a blackish central abdominal sclerite; they fly to tubuliferous Compositae (Cir-
sium, Carduus, Cynara, Arctium). Males and gynoparae return to Elaeagnus
in autumn. Widely distributed in temperate and warm temperate regions of
the world. See also Hille Ris Lambers, 1953a; B & E, 1984, p. 249. 2n = 16.

Capitophorus himalayensis A.K. Ghosh, M.R. Ghosh and Raychaudhuri

Apterae slender, BL 1.4-1.6 mm, pale except for brownish frons and dusky
tips to tarsi and siphunculi. Alatae and other morphs are undescribed and the
life cycle is unknown. On Hippophae sp. (Saha and Chakrabarti, 1988d) in
northwest India (Uttar Pradesh, West Bengal).

Capitophorus hippophaes (Walker) Apterae in spring colonies on Elaeagna-

ceae (Elaeagnus spp., Hippophae spp.) are pale green, slender, with a faint
pattern of green spots; BL 1.5-2.1 mm. Fundatrices are very different; broadly
oval, greenish with reddish spots and their antennae are dark, 5-segmented and
have a short processus terminalis. Alatae, produced in the second and third
generations, are greyish-green with a black head and thorax, dark antennae,
legs and siphunculi and a large quadrate dark green patch on the dorsal
abdomen; they migrate to various Polygonum spp. (Hille Ris Lambers,
1953a). Capitophorus hippophaes sensu stricto occurs in Europe, southwest
Asia, and introduced into North America. Populations on Elaeagnus spp. and
Polygonaceae in the Far East differ in the absence of any conspicuous capitate
hairs on ANT I and ABD TERG 6 and are usually regarded as a subspecies,
C. hippophaes javanicus; this form occurs on Polygonaceae in Southeast
Asia, Australia, New Zealand, South America and also in California, USA
(Corpuz-Raros and Cook, 1974). 2n = 10 (for both forms).

Capitophorus meghalayensis Basu and Raychaudhuri Apterae pale, narrow-

bodied; BL about 1.3-1.5 mm. Alatae with a faint brownish patch extending
from ABD TERG 2-6 (R.C. Basu and Raychaudhuri, 1976b); apparently there
is no clearly defined central sclerite. On Elaeagnus sp. in Meghalaya, India.
Oviparae are described, but the life cycle and secondary hosts, if any, are
unknown.

Capitophorus pakansus Hottes and Frison (= vandergooti HRL) Apterae

on Elaeagnaceae in spring are pale, narrow-bodied; BL 1.6-2.Omm. Fun-
datrices are oval and mottled dark green. Alatae have black head and thorax
and a quadrate dorsal abdominal patch; they fly for the summer to lnula spp.
Primary hosts are Elaeagnus spp. (argentea, commutata) and sometimes Hip-
pophaes rhamnoides (Norway; BMNH colln), although gynoparae in Holland
did not produce oviparae on Hippophae (Hille Ris Lambers, 1953a). In
northwest Europe and eastern North America. 2n = 16*.
594 Carolinaia

Capitophorus shepherdiae Gillette and Bragg Apterae are pale, spindle-

shaped, BL 1.4-1.7 mm, in spring colonies on Shepherdia argentea, which is
the only authenticated primary host. Probably there is host alternation to
Ambrosia spp., although this needs confirmation with transference tests
(Corpuz-Raros and Cook, 1974). In western USA (California, Colorado,
Utah).

Capitophorus similis van der Goot Apterae in spring colonies on Elaeagn-

aceae are pale green, spindle-shaped, BL 1.7-2.3 mm. Fundatrices are more
oval in shape with 5-segmented antennae. Alatae have a creamy-coloured
abdomen with a dark quadrate dorsal patch, and migrate to Tussilago spp.
and Petasites spp. Males and gynoparae migrate back to Hippophae or
Elaeagnus in September-October (Hille Ris Lambers, 1953a). Loher and
Lampel (1983) studied the factors controlling sexual morph production in this
species. In Europe and Central Asia; anholocyclic overwintering populations
may occur on secondary hosts in warmer climates (Patti, 1983).

Capitophorus xanthii (Oestlund) Apterae on Elaeagnaceae in spring have

not been described. The summer generations occur on Xanthium spp. and
males and oviparae were collected on Hippophae rhamnoides in Colorado in
August-October (Palmer, 1952).

CAROLINAIA Wilson Aphidinae: Macrosiphini

About 17 species typically associated with Rhus, or with host alternation bet-
ween Rhus and Cyperaceae. Remaudière and Muñoz Viveros (1992) recently
expanded the concept of the genus to include Glabromyzus (four North
American spp.) and Juncomyzus (five East Asian spp.). Carolinaia sensu
stricto and all except one species of Glabromyzus are associated only with
Rhus subgenus Toxicodendron, so they do not concern us here.

Carolinaia (Juncomyzus) floris (Miyazaki) Appearance in life on Rhus

not recorded; on secondary hosts, apterae are yellowish to reddish-brown.
Described from Juncus and Carex (Miyazaki, 1971), but fundatrices and
alatae collected on Rhus ambigua in Hokkaido, Japan (BMNH colln, leg. M.
Miyazaki) were subsequently identified as this species. An alata trapped in
Korea also seems to be C. (J.) floris (BMNH colln, leg. W. Paik). BL of
fundatrices 2.1-2.2 mm. Biology not studied; the host alternation requires
experimental confirmation.

Carolinaia (Juncomyzus) japonicus (Takahashi) Apterae shining yellowish-

brown, with black antennae, femora, tibial apices, siphunculi and CAUDA;
BL c. 1 .9 mm. On Rhus trichocarpa and Rh. verniciflua in Japan (Takahashi,
 Cavariella 595

1924a, as Myzus rhois; Takahashi, 1963a, as Sitomyzus japonicus - but the

fundatrix described under this name is C. (J.) nigra, see Miyazaki, 1971). Pro-
bably heteroecious, but secondary host unknown; oviparae on Rh. trichocarpa
in September (Remaudière and Mufioz Viveros, 1992).

Carolinaia (Juncomyzus) nigra (Miyazaki) Apterae elongate oval, blackish-

brown; BL 1.8-2.0 mm. On Rhus trichocarpa in Japan (Miyazaki, 1971).
Probably heteroecious, but secondary host unknown; oviparae, an alate
gynopara and an alate male were collected on Rh. trichocarpa in September
(Remaudière and Mufloz Viveros, 1992).

Carolinaia (Glabromyzus) rhois (Monell) Apterae are yellow-brown with

black clavate siphunculi; BL 1.9-2.3 mm. On undersides of leaves of Rhus
glabra and Rh. typhina in North America. Alatae have a brownish-yellow or
greenish-yellow abdomen with dusky dorsal markings. Monoecious holocyclic
on Rhus, with alate males. There is possibly a partial migration to Gramineae,
but most grass-feeding populations of Carolinaia (Glabromyzus) are probably
G. howardii (Hille Ris Lambers, 1966d; B & E, 1984, p. 278).

CASTANOCALLIS Zhang and Zhong

Drepanosiphinae: Phyllaphidini

Three species, all recorded only from Castanea bungeana in China. All
viviparae are alate. The genus resembles Myzocallis (Nippocallis) in wing
pigmentation, presence of spinal and marginal tubercles and a rather short
processus terminalis, but the species of Castanocallis are much more robust.
See Zhang and Zhong (1981c) for further information. Castanocallis
castanocallis has 2n = 14*.

CAVARIELLA del Guercio Aphidinae: Macrosiphini

A holarctic genus with a characteristic supracaudal process, containing around

30 species, about half of them in Asia. Most species host-alternate between
Salix and Umbelliferae or the closely-related Araliaceae; one is known to
be monoecious on Salix and several others are recorded only from Salix
with life cycle unknown. Parasitoids are Aphidius salicis (ref: Stary, 1973)
and Ephedrus salicicola (ref: D. Raychaudhuri, 1990). Taxonomie revisions
and/or keys are available for Europe (Hille Ris Lambers, 1947b), Fenno-
scandia and Denmark (Heie, 1992), Siberia (Ivanoskaya, 1980), eastern Siberia
596 Cavariella

(Pashchenko, 1988b), Japan (Miyazaki, 1971), India (Raychaudhuri, 1980)

and North America (Stroyan, 1969).
Cavariella aegopodii (Scopoli) Willow-Carrot Aphid Apterae in spring
colonies on young leaves and catkins of Salix spp. are oval, somewhat dor-
soventrally flattened, pale green, greenish-yellow or yellowish-white; BL
1.5-2.0 mm. Alatae produced from the second generation onwards have black
head and thorax and a dark green dorsal abdominal patch, mostly migrating
to wild and cultivated Umbelliferae in May-June, but sometimes recolonizing
willow, so that populations can persist on willow into July and August (Dunn,
1965). The return migration from Umbelliferae to willow occurs in late
September to early November. Preferred primary hosts seem to be S. fragilis
and S. alba; some Salix spp. seem only to be colonized in spring, by alatae
from overwintered anholocyclic populations, and are thus acting as secondary
hosts. Cavariella aegopodii is widespread throughout temperate and warm
temperate parts of the world (B & E, 1984, p. 250). 2n = 10.
Cavariella (Cavaraiellia) aquatica Gillette and Bragg (= hillerislambersi
Ossiannilsson) Apterae in spring colonies on Salix spp. (especially lapponum,
nigricans, purpurea, urbaniana) are narrow-bodied, green, BL 1.4-2.0 mm.
Fundatrices are remarkable for this genus in their complete lack of siphunculi
(Stroyan, 1969). Alatae have blackish head and thorax and dark dorsal abdom-
inal bands sometimes partly fused into a solid patch; they migrate to found
summer colonies on plants growing in water or marshy situations. Unusually
for this genus, a wide variety of secondary hosts including Gramineae
and Juncaceae may be colonized. Cavariella aquatica is recorded, often at
high altitudes, from Europe, Iran, Afghanistan, India, Pakistan, Siberia
(Kamchatka, Sakhalin, Primorskiya Kray), USA (Colorado) and Canada
(Northwest Territories; BMNH colln, leg. R. O'Doherty). See also Palmer
(1952) as Aspidaphis aquatica and Stroyan (1964a) as C. hillerislambersi.
2n = 8*.
Cavariella araliae Takahashi Presumably host-alternating between Salix
spp. and Araliaceae (Aralia, Schieffera, Tetrapanax), but this needs to be con-
firmed, as no fundatrices or sexual morphs have been described and the only
Salix host identified to species is S. warburgii, in Taiwan, on which popula-
tions overwinter anholocyclically along the mid-ribs of the leaves (Takahashi,
1923; as neocapreae). Recorded from China, Japan, Korea, Okinawa and east
Siberia. It seems possible that the aphids from Salix in India described as
C. biswasi (q.v.) are the primary host forms of araliae. 2n = 14.
Cavariella archangelicae (Scopoli) Apterae in spring colonies on Salix
spp. are pale yellow-green to straw-coloured, BL 2.0-2.3 mm. Alatae have
black head and thorax and broad dark green cross-bands on the dorsal
abdomen, sometimes merged into a solid patch; they migrate to Angelica
spp. (including Archangelica) from late May to July. Stroyan (1964a) discusses
differences between archangelicae and the very similar C. konoi. In northwest
Europe, including Iceland (B & E, 1984, p. 251). 2n = 6.
 Cavariella 597

Cavariella aspidaphoides Hille Ris Lambers Apterae on Salix in spring are

green, elongate pear-shaped with pointed abdomen; BL 1.7-2.2 mm. Often
they form mixed colonies with C. aegopodii. Alatae migrate in May to
Umbelliferae; the only recorded host is Daucus maximus. Only known from
Israel and Iran (Hille Ris Lambers, 1970a).

Cavariella biswasi Ghosh, Basu and Raychaudhuri Apterae in spring on

Salix spp. (incl. babylonica, elegans) are pale (green?), spindle-shaped, BL
1.7-1.9 mm. Alatae, produced in July, have broad dark partially coalesced
bands on ABD TERG 3-5, and smaller sclerites on other tergites. The secon-
dary host is unknown, but this may be the primary host form of C. araliae
(q.v.), from which it differs only in the size and form of the process on ABD
TERG 8 (A.K. Ghosh et al., 1969a).

Cavariella borealis Hille Ris Lambers Monoecious holocyclic on Salix arctica

in northwest Greenland and northern Canada (Northwest Territories), with an
abbreviated cycle, producing oviparae and apterous and/or alate males in
July. Colour of apterae in life not recorded; they are very variable in size and
form, with BL 0.7-2.3 mm. Alatae can be brachypterous, intermediate or
fully-winged (Hille Ris Lambers, 1960b). 2n = 6*.

Cavariella digitata Hille Ris Lambers Appearance in life unknown, apterae

probably pale yellowish or greenish; spindle-shaped, rather flattened, BL
1.7-2.3 mm. Alatae have a dusky, somewhat perforated, dorsal abdominal
sclerotic patch. On unidentified Salix sp(p)., only recorded from Utah, USA
(Hille Ris Lambers, 1970a). Life cycle is unclear; populations persist on Salix
into August, suggesting that if host alternation occurs at all then the migration
must be incomplete. Alate males occur in October.

Cavariella hendersoni Knowlton and Smith Appearance in life unknown.

On leaves of unidentified Salix sp. in Utah, USA. Life cycle is unknown. The
original description was of alatae only, which resemble those of C. salicis
(Monell) in having secondary rhinaria on ANT IV and dorsal abdominal cross-
bands separate or only partially coalesced. BL of alate paratypes 1.7-1.9 mm
(Palmer, 1952). Differences from salicis include the usual presence of acces-
sory hairs on R IV+V and more slender siphunculi. Stroyan (1969) described
some characters of presumed apterae of this species from Logan Canyon,
Utah, and its identification in the key is based on these specimens, now in the
BMNH collection. There are records of hendersoni from seven other states
of USA and from Ontario, Canada (Smith and Parron, 1978), but most of
these are likely to be misidentifications (see Stroyan, 1969).

Cavariella indica Maity and Chakrabarti Appearance in life unknown,

probably pale green; BL of aptera c. 1.9-2.1mm. Alata is undescribed. In
colonies on upper surfaces of leaves and on apical growth of Salix babylonica
in Uttar Pradesh, India (Maity et al., 1982). Monoecious holocyclic, with
oviparae and males in October-December (Medda et al., 1990). On the basis
598 Cavariella

of the published description, indica seems morphologically very similar to

C. aegopodii, except that R IV+V of the aptera is reported to have 2 accessory
hairs.

Cavariella intermedia Hille Ris Lambers Apterae are evenly light green,
rather broadly spindle-shaped, BL 1.4-2.3 mm. Alatae differ from aegopodii
by having only a rather indistinct smoky dorsal abdominal patch due to pale,
partly coalescent sclerotic bars on ABD TERG 3-6 (becoming somewhat
darker in alatae developing at lower temperatures). Alatae also differ from
aegopodii by having 2-7 secondary rhinaria on the fourth antennal segment.
Recorded from Salix aurita, S. arbuscula and S. phylicifolia in Belgium,
Netherlands, Switzerland and UK. Life cycle is not completely known; Hille
Ris Lambers (1970a) found apterae producing oviparae and alatoid nymphs
(presumed males) on Salix in September, which seems to rule out host alterna-
tion. 2n = 6.

Cavariella japonica (Essig and Kuwana) Apterae are dusky yellowish,

pinkish, purplish or greyish-brown, with dark distal sections to antennae, legs
and siphunculi; BL 1.5-2.4 mm. Alatae have a dark orange-yellow abdomen
with a black dorsal patch. On Salix spp. in Japan, Korea (BMNH colln) and
Siberia (Kamchatka, Primorskiya Kray; Pashchenko, 1988b), migrating for
the summer to Umbelliferae (Angelica, Anthriscus, Cryptotaenia, Distaenia,
Torilis; Miyazaki, 1971). Takahashi (1937b) also recorded a wholly black form
of aptera from high altitudes in Taiwan, on S. fulvipubescens. Sexuales are
undescribed. 2n = 8.

Cavariella konoi Takahashi Apterae in spring colonies on Salix are green

or yellowish-green, often with two darker green longitudinal stripes; BL
c. 2.3-2.5 mm. Alatae have a solid dark dorsal abdominal patch on ABD
TERG 3-5 and two characteristic small roundish spots close together on ABD
TERG 6, posterior to the level of the bases of the siphunculi. Other differences
from the biologically and morphologically similar C. archangelicae were
discussed by Stroyan (1964a), with descriptions also of males and oviparae.
Recorded from Europe (incl. Iceland), North America, Mexico, Japan, China,
Korea and Siberia. Host-alternating between Salix spp. and Umbelliferae; in
Europe the only secondary hosts are Angelica spp., whereas in Japan
Heracleum is also colonized (Miyazaki, 1971), and in America secondary host
populations are recorded on Angelica, Apium, Cicuta and Sium (Stroyan,
1969). 2n = 8.

Cavariella nigra Basu Apterae in spring colonies on Salix are dusky black
including antennae, legs, siphunculi and CAUDA, pear-shaped, somewhat
flattened; BL 1.9-2.5 mm. Alatae have a dark brown central dorsal abdominal
patch. The black coloration of the aptera is unusual for a Cavariella; the dark
form of japonica reported from Taiwan is possibly this species, although
japonica from Japan has a longer last rostral segment (R IV+V 1.4-1.6 x HT
II, as opposed to R IV+V 1.0-1.2 x HT II in nigra). Cavariella nigra is
 Cavariella 599

also unusual in having 0-1 secondary rhinaria near the middle of ANT V of
the aptera (none on III or IV). On undersides of leaves of Salix elegans in West
Bengal, India (A.N. Basu, 1964). The life cycle is unknown.

Cavariella nipponica Takahashi Apterae are whitish; BL about 1.6mm.

Alatae have dark dorsal abdominal cross-bands fused into a solid patch on
tergites 3-5. Apparently host-alternating in China, Japan, Korea and Siberia
(incl. Kamchatka) between Salix spp. and Umbelliferae (Angelica, Heracleum,
Pachypleurum), although the life cycle does not appear to have been studied
and no sexuales are described (Takahashi, 1961c; Miyazaki, 1971; Pash-
chenko, 1988b).

Cavariella pastinacae (Linnaeus) (= essigi Gillette and Bragg) Apterae in

spring colonies on leaves and new growth of Salix spp. are rather shiny light
green, with antennae mainly pale (darker on base VI) and legs pale except
for tarsi; BL 1.6-2.2 mm. Alatae have very broad dark bands on ABD TERG
3-5 coalescing to form an almost solid dark green to black trapezoid patch.
Host alternation occurs mainly to Heracleum, less commonly to Pastinaca.
Oviparae and males are found on Salix in October. Cavariella pastinacae
occurs throughout Europe and is also widely distributed in North America.
2n = 8.

Cavariella pustula Essig Apterae pale greenish-yellow to orange, with

siphunculi and distal extremities of antennae and legs dusky; BL about
1.6 mm. Alatae have black head and thorax and pale greenish-yellow abdomen
with two light green dorsal longitudinal stripes (Essig, 1937). On new growth
and undersides of leaves of Salix agrophylla (BMNH colln; leg. R.C. Dickson)
in California, and also recorded from unidentified Salix in California, Utah
and British Columbia (BMNH colln). Life cycle unknown.

Cavariella rutila Mamontova Apterae in spring colonies on leaves and new

growth of Salix are reddish or bright yellowish with red markings; BL
c. 1.8-1.9 mm. Alatae are reddish-yellow with broad dark brown cross-bands
on dorsal abdomen (Mamontova-Solukha, 1961). Only recorded from Salix
purpurea in Ukraine. Life cycle is unknown.

Cavariella salicicola Matsumura (= bicaudata Essig and Kuwana) Apterae

in spring colonies on new growth of Salix are yellowish-green to green with
bluish pruinosity (Moritsu, 1983); BL c. 2.0-2.2 mm. Immature alatae are
bluish-grey with paired dark segmental markings, adult alatae have black head
and thorax and variably developed dark dorsal abdominal cross-bands or
pleural markings. Recorded from numerous Salix spp. and also from Chosenia
arbustifolia in East Asia (India, Japan, Korea, China, Tibet, Thailand and
east Siberia) and probably also in southwest Asia and Eastern Europe (see
below). Migration occurs in April-May to various Umbelliferae (incl. Apium,
Cryptotaenia, Oenanthe, Levisticum, Sanicula, Sium), with a preference for
600 Cerataphis

plants growing in water or marshy situations. Sexuales are found on Salix in

late October (Wang et al., 1988).
This species closely resembles C. cicutae (Koch), which occurs on aquatic
Umbelliferae in Europe (Hille Ris Lambers, 1947b). There is some confusion
over the life cycle of cicutae, which has been reported to host-alternate
between Salix and Umbelliferae in the Ukraine (Mamontova-Solukha, 1961)
and Iran (Hodjat, 1984). However, oviparae are produced by cicutae in
October on Sium latifolium, indicating that it is morioecius holocyclic on
aquatic Umbelliferae (Hille Ris Lambers, 1952). Specimens from Iran in the
BMNH collection, previously identified as cicutae, are indistinguishable from
East Asian salicicola, so all host-alternating populations are here considered
to be the latter species. 2n = 10 (Chen and Zhang, 1985b).

Cavariella salicis (Monell) Apterae on Salix, according to Monell's original

notes, are green with two darker green longitudinal stripes on the abdomen
(Stroyan, 1969); BL 1.6-2.1mm. Alatae have dusky head and thorax and
green abdomen with irregular darker green markings. On Salix spp. in eastern
and central USA, with at least a partial migration to Umbelliferae (Angelica,
Sium). Stroyan (1969) clarified the status of salicis which was for many years
confused with aegopodii and other species.

Cavariella takahashii Hille Ris Lambers Only the alata of this species is
described, so it could not be included in the key to Cavariella on Salix. BL
of alata about 2.2-2.3 mm; dorsal abdominal markings are indistinct and the
femora have numerous long wavy hairs as well as a number of much shorter
hairs on distal half similar to those on tibiae. The antennae have a short pro-
cessus terminalis and numerous secondary rhinaria; 51-52 on ANT III, 10-11
on IV and 3-5 on V. On Salix spp. in April in Japan (Hille Ris Lambers,
1965). Appearance in life and biology are unknown.

Cavariella theobaldi (Gillette and Bragg) Apterae in spring colonies on

leaves and new growth of Salix spp. are yellowish-green to green; BL
1.6-2.0 mm. Alatae have variably-developed dark dorsal abdominal cross-
bands, often forming a central trapezoid patch. They fly in late April-June
to Umbelliferae (Heracleum, Pastinaca, or more rarely Aegopodium,
Angelica, Chaerophyllum). The return migration occurs in late September-
October. In Europe, southwest Asia, Siberia (Pashchenko, 1988b), and widely
distributed in North America. 2n = 8.

CERATAPHIS Lichtenstein Hormaphidinae: Cerataphidini

About six Southeast Asian species, some of which may host-alternate between
galls on Styrax and monocotyledonous secondary hosts (Araceae, bamboos,
 Ceratoglyphina 601

Orchidaceae, Palmae, Pandanaceae) in their native regions, where their

biology is largely unstudied. They have unusual, yeast-like extracellular
symbionts, found also in Tuberaphis, Glyphinaphis and Hamiltonaphis,
indicating a monophyletic relationship between these genera (Aoki et al.,
1993). Apterae on secondary hosts are dark, round, scale-like, with a marginal
ring of wax. Alatae have a once-branched media and 5-segmented antennae.
Noordam (1991) provided a generic diagnosis and account of Javanese species.
For the palm-feeding species (variabilis, lataniae, formosana) see B & E, 1984,
p. 252.

Cerataphis bambusifoliae Takahashi Apterae on stems and undersides of

leaves of bamboos (Bambusa, Arundinaria, Pleioblastus) are black, aleyrodi-
form, with a white wax fringe; BL 1.5-2.5 mm. Alatae have antennae with
annular secondary rhinaria distributed III 24-34, IV 9-13, V 8-13. Life
cycle is unstudied; alatae (sexuparae?) are produced in large numbers in
September-October (Takahashi, 1939a; Saha and Chakrabarti, 1988b), but
their destination is unknown. Recorded from China, Taiwan and Uttar
Pradesh, India.

Cerataphis fransseni (Hille Ris Lambers) (= C. variabilis Hille Ris Lambers),

Plate 3a The galls of this species on Styrax are bag-like, greyish-green,
10-47 mm long and 5-14 mm wide, with an apical slit-shaped opening. They
arise from the stem, either terminally or in a leaf axil, and are produced at
any time of year (Stern et al., 1994). Apterae in the galls are small (BL about
1.3mm) with spine-like frontal hairs including one pair of particularly large
ones, short 5-segmented antennae and siphuncular pores on small cones with
a ring of 3-5 hairs. Second instar soldier larvae move in and around the galls
(Noordam, 1991). Alatae emerging from galls have antennal secondary
rhinaria distributed III 15-23, IV 2-8, V 0-4, and BL 1.2-1.6 mm. They fly
to found colonies on various species of palms; for a long time the palm-feeding
generations have been known as C. palmae or C. variabilis, which must now
be regarded as synonyms (Stern et al., 1994). On Styrax benzoin in Java (Hille
Ris Lambers, 1933a; Noordam, 1991) and Malaya (Takahashi, 1950; as ssp.
kepongensis, now regarded as a synonym). Life cycle unknown.

CERATOGLYPHINA van der Goot

Hormaphidinae: Cerataphidini

Aphids similar to Astegopteryx but distinguishable from that genus and most
other Hormaphidinae by the rounded CAUDA and anal plate. The primary
host has recently been shown to be Styrax as in other Cerataphidini, but many
populations are probably anholocyclic on bamboos. Noordam (1991) reviewed
the species in Java.
602 Ceratovacuna

Ceratoglyphina bambusae van der Goot Apterae on bamboos are oval-

bodied, dark brown or dark marbled greenish-black, with pale brown legs, and
a flat fringe of white wax; BL 1.3-2.2 mm. Immatures are yellowish to green,
with brown heads in later instars (Noordam, 1991). Dense colonies form near
bases of leaves, often attended by ants. Collected on bamboos (Bambusa,
Dendrocalamus, Gigantochloa, Phyllostachys, Schizostachyum) in China,
Indonesia and Malaysia. 2n = 12 (Chen and Zhang, 1985b).

Ceratoglyphina bengalensis L.K. Ghosh Apterae are pale brown with two
rather indistinct longitudinal green stripes and a narrow, flat, wax fringe;
BL 2.1-2.5 mm. On undetermined bamboos in India (West Bengal, Assam;
M.R. Ghosh et al., 1977b) and also recorded from Java, where the aphids were
found living on the border of the sheath and at the leaf-bases of young shoots
(Noordam, 1991).

Ceratoglyphina styracicola (Takahashi) The galls on Styrax suberifolia in

Taiwan are large, almost globular, somewhat like a compact cauliflower head,
55-80 mm in diameter, in clusters arising from the stem (Fig. 124J, p. 573; and
Kurosu and Aoki, 1991 a, as C. bambusae). The surface is covered with white
wax powder, and the inside is a labyrinth of green plant tissue (Takahashi,
1924a; Aoki et al., 1977). Gall inhabitants, which may number 200,000 per
gall, include yellowish-brown adult apterae, immatures developing normally,
and strongly sclerotized 'biters', which do not usually undergo development
beyond the second instar (Aoki, 1979a; Kurosu and Aoki, 1991b, as C. bam-
busae). Aphids readily wander from these galls and may invade galls of
Astegopteryx bambucifoliae on the same plant (Kurosu and Aoki, 1990a).
Alatae emerge from late December to March and migrate to bamboos
(Bambusa, Pleioblastus); they have secondary rhinaria distributed III 17-23,
IV 6-9, V 4-8. [Aoki (pers. comm.) has informed us that the synonymy with
C. bambusae proposed by Aoki and Kurosu (1989) is incorrect, as the bamboo-
feeding generations in Taiwan show differences from true bambusae.]

CERATOVACUNA Zehntner Hormaphidinae: Cerataphidini

A genus in East and Southeast Asia, closely related to Astegopteryx and with
a similar alternation between Styrax and Gramineae, although this has only
been confirmed for one species. An indefinite number of species form dense
colonies on the undersides of bamboo leaves. The apterae are small to
medium-sized, yellow to brown in colour, and secrete much wax. Alatae of
the bamboo feeders are rare or unknown. As in the case of Astegopteryx,
characters used to separate species are frequently those known to vary accord-
ing to the environment, such as the degree of development of the wax glands,
and the genus is in need of further taxonomie revision. Accounts are available
 Ceratovacuna 603

for Japan (Takahashi, 1958c), Java (Noordam, 1991), Korea (Paik, 1965),
India (M.R. Ghosh et al., 1977b) and Taiwan (Liao, 1976).

Ceratovacuna floccifera Noordam Appearance in life unknown, but apterae

presumably with flocculent wax; BL 1.4-1.6 mm. First instar soldier larvae
occur with enlarged fore legs and very long frontal horns. On undetermined
bamboos in Java (Noordam, 1991). Life cycle is unknown. Closely related to
and possibly synonymous with C. japonica or C. longifila.

Ceratovacuna hoffmanni (Takahashi) Apterae of unknown appearance in

life but probably darkish and with much wax secretion; BL 1.8-2.7 mm. On
bamboos (Bambusa, Schizostachyum) in China and Sumatra. Indoregma
bambusae Chakrabarti and Maity, 1982, described from northwest Himalaya
on Bambusa sp., is possibly the same species. Ceratovacuna longifila and
C. japonica (see below) are also similar aphids, differing mainly in degrees
of wax gland development; further work is needed to show if any or all of
them are distinct species.

Ceratovacuna indica M.R. Ghosh, Pal and Raychaudhuri Apterae pear-

shaped, yellowish in life and covered with white wax; BL 1.4-1.7 mm. In
colonies on the undersides of older leaves of unidentified bamboos in West
Bengal (M.R. Ghosh et al., 1977a). Possibly a synonym of Indoregma
bambusae (q.v.).

Ceratovacuna japonica (Takahashi) Apterae on bamboos are pinkish-

brown to dark brown in life, secreting white wax tufts or filaments; BL
1.5-2.0 mm. On undersides of leaves of Bambusa spp. and Pleioblastus chino
in Taiwan, Japan and Korea. Anholocyclic populations live throughout the
year on bamboos, but there is also host alternation to Styrax japonica. Galls
are like those of C. nekoashi, but have a longitudinal suture (Kurosu et al.,
1990, as C. pseudostyracophila; Aoki and Kurosu, 199la). Two other species
described from bamboos in Taiwan, brevicornis Takahashi and longifila
Takahashi (the latter secreting very long wax filaments), may be synonyms of
japonica (but see Liao, 1976). 2n=12.

Ceratovacuna keduemis Noordam Apterae dull black, with paired spinal

and marginal rows of short, conical wax tufts; BL 1.6-1.8 mm. On undersides
of leaves of undetermined bamboos, often forming large colonies, in Java
(Noordam, 1991). Life cycle unknown. Possibly a synonym of C. hoffmanni.

Ceratovacuna nekoashi (Sasaki) The gall on Styrax in spring resembles a

bunch of bananas (or a cat's paw, which is 'nekoashi' in Japanese) and is
developed from an axillary bud (Fig. 124K, p. 573; Kurosu & Aoki, 1990b).
Alate emigrants (BL c. 1.7-1.8 mm, with dark cross-bands on ABD TERG
6-8) emerge in June-July and migrate to Gramineae (Apluda, Arthraxon,
Microstegium, Miscanthus, Oplismenus, Panicum). Sexuparae returning to
Styrax in October differ from the spring emigrants by having frontal horns
604 Cervaphis

and less sclerotization of ABD TERG 6 (Takahashi, 1958c). They deposit first
instar sexuales on the leaves; these move to the branches without moulting and
develop to maturity without feeding, the eggs being laid on bark (Kurosu and
Aoki, 1990c). In Japan, Taiwan, Korea and Uttar Pradesh, India (Chakrabarti
and Maity, 1982). 2n = 12.

Ceratovacuna sylvestrii (Takahashi) Apterae yellow to reddish-brown or

almost black in life, with numerous white wax tufts over dorsum and sides
of body; BL 1.6-2.3 mm. On undersides of leaves, near their bases, of both
Arundinaria spp. and Bambusa spp. Recorded from Taiwan and India (as
subglandulosa Hille Ris Lambers and Basu, 1966). Agarwala et al. (1987)
described a parasitoid. 2n = 12 (Khuda-Bukhsh and Kar, 1987).

CERVAPHIS van der Goot Greenideinae: Cervaphidini

A genus with five species in India and Southeast Asia, in which the apterae
have remarkable branched marginal hair-bearing processes. In alatae these
processes are reduced to low, flat hair-bearing tubercles, except on the head.
The siphunculi are long, cylindrical, slightly curved outwards and a little
swollen subapically, where there is a ring of small hairs. For revisions of
Cervaphis see Hille Ris Lambers (1956a) and A.K. Ghosh (1982a).

Cervaphis quercus Takahashi Apterae are yellow with pale siphunculi, BL

about 1.7mm, feeding on undersides of leaves and new growth of Quercus
spp. in East Asia (recorded from Japan, Taiwan, Thailand, China, Korea and
Assam, India). Sometimes ant-attended. Alatae have brownish head, thorax
and dorsal abdominal markings including a large central patch and siphunculi
almost black (Takahashi, 1924a). Biology is largely unstudied; alate oviparae
have been found in Japan from July onwards, produced by apterous
sexuparae, but males have not been recorded (Takahashi, 1923). 2n = 8.

Cervaphis schouteniae van der Goot (= cambodiensis Takahashi) Apterae

are whitish-green to yellowish-white with pale yellowish-brown siphunculi;
BL 1.5-1.9 mm. Alatae have brownish-yellow head and thorax, pale yellow
abdomen, dark antennae and black siphunculi. On undersides of young
leaves, feeding along main veins near petioles. Host plants include trees and
shrubs mainly in Tiliaceae (Actinophora, Grewia, Microcos, Schoutenia), but
also recorded from Pterospermum (Sterculiaceae), Dipterocarpus (Diptero-
carpaceae) and Zizyphus (Rhamnaceae). Recorded from Malaysia, Indonesia,
Philippines, Thailand, Cambodia and India. Sexuales are unrecorded. Popula-
tion trends in northeast India were studied by Agarwala and Dixon (1986),
developmental stages are described by Agarwala et al. (1985). Parasitoid:
Trioxys peniculatus (Agarwala et al., 1987).
 Chaitophorus 605

CHAETOGEOICA Tao Pemphiginae: Fordini

A genus of three species, two of which are only known from grass roots in
India. They are related to Geoica but the emigrant alatae have ciliated secon-
dary rhinaria and their embryos have fewer dorsal hairs.

Chaetogeoica foliodentata (Tao) The gall on Pistacia sinensis is a green and

pinkish elongate sac about 4 cm long with a terminal opening, arising from
a leaflet (Fig. 126D). Emigrant alatae, BL 1.4-1.9 mm, are mature in May
(Remaudiére and Tao, 1958). The secondary hosts are unknown, but pre-
sumably roots of Gramineae. Recorded from Pistacia in China and Taiwan,
and single alatae of Chaetogeoica have also been trapped or collected in
Korea (Paik, 1972), the Philippines, Nepal (BMNH colln) and Sikkim, India
(A.K. Ghosh and Raychaudhuri, 1968a).

CHAITOPHORUS Koch Chaitophorinae: Chaitophorini

A large holarctic genus (about 88 species) associated exclusively with Salica-

ceae, with individual species feeding on either Populus or Salix but never both,
and often with a high degree of host specificity. They are very hairy, small
to medium-sized aphids with short, stump-shaped siphunculi having at least
some trace of reticulate ornamentation. The dorsal cuticle of the aptera is
usually more-or-less completely sclerotic and variably sculptured over at least
ABD TERG 2-6. Alatae often have dark segmental cross-bands and marginal
sclerites. The CAUDA is usually knobbed, but (unlike most Drepanosiphinae
with knobbed CAUDAS) the anal plate is entire. Pintera (1987) fully revised
the palaearctic fauna (58 species) and for North America there are revisions
by Hille Ris Lambers (1960a) and Richards (1972a - Canada only). The para-
sitoids Lysiphlebus salicaphis (refs: Smith, 1944; Starý, 1976) and Aphelinus
aureus (Gahan, 1924) specialize on Chaitophorus. Aphelinus fulvus also
parasitizes Pterocomma spp. on the same hosts (Kalina and Starý, 1976).

Chaitophorus capreae (Mosley) Apterae white to yellowish-white; BL 0.8-

1.9mm. Alatae without distinct dark dorsal abdominal markings. On broad-
leaved Salix spp., scattered on undersides of leaves, not attended by ants (see
Heie, 1982; Pintera, 1987). Apterous males (yellow) and oviparae occur in
October-November. Widespread in Europe, and eastward to Central Asia.
2n = 30. Chaitophorus flavissimus Mamontova, described from Salix caprea
in Ukraine, is very similar to capreae but has a less constricted CAUDA and
possibly longer antennal hairs.
606 Chaitophorus

Chaitophorus clarus Tseng and Tao Appearance of aptera in life unknown,

probably pale; BL about 1.3-1.4 mm. Alata withut any distinct dorsal abdomi-
nal markings. On Populus simonii in China (see Pintera, 1987). Biology
unknown.

Chaitophorus crinitus Ivanoskaya Apterae greenish-brown; BL 1.8-2.0 mm.

On leaves of unidentified Salix sp., attended by ants, in Siberia (see Pintera,
1987).

Chaitophorus crucis (Essig) Apterae according to the original description

(Essig, 1912a) have distinctive pigmentation in life; rich green with broad
light yellow-green spinal stripe from the base of the head to the CAUDA,
and a transverse band of similar width and colour across the anterior abdom-
inal tergites, making a large cross. BL about 1.3mm. Alatae have dark
dorsal abdominal markings. Found in dense colonies on undersides of leaves
of Salix macrostachyae in California, but not recorded since the original
description.

Chaitophorus diversisetosus Szelegiewicz and Czylok Apterae according to

the original description (Szelegiewicz and Czylok, 1981) are blackish-green
with a paler central patch, and have dark legs and tips to the siphunculi; BL
about 1.8 mm. Alatae have dark dorsal abdominal cross-bands. In ant-attended
colonies on undersides of leaves of Salix acutifolia in Poland. A related form,
Ch. diversisetosus ssp. austriacus, was described from Salix purpurea in
Austria, France and Switzerland (Pintera, 1987). Apterae of this subspecies
are 'very pale green with a bright green lyre pattern and short median stripe
sometimes darkened', with legs also very dark; BL 1.3-1.6 mm. Colonies, on
upper as well as undersides of leaves, were not attended by ants. An apterous
male was collected in August.

Chaitophorus dorocolus Matsumura Apterae elongate-bodied, presumably

pale-coloured and, according to original description, covered with a woolly
wax secretion (an unusual feature for a Chaitophorus); BL about 1.3mm.
Alatae identified as this species by Pintera (1987) have broad dark dorsal
abdominal cross-bands, and BL 1.8-2.1mm. On undersides of leaves of
Populus sieboldii in Hokkaido, Japan, and also reported to occur on P. davi-
diana in east Siberia (Pashchenko, 1988b) and Populus sp. in West Bengal,
India (Maity and Chakrabarti, 1981). Higuchi's (1972) account of dorocolus,
and probably most records from P. maximowiczii, refer to C. inouye.
2n = 14.

Chaitophorus eoessigi Hille Ris Lambers Apterae variable in colour and

markings even within colonies, often with a dark dorsal lyre-shaped pattern,
or almost wholly dark except for a pale spinal stripe; BL 1.1-1.9 mm. Alatae
have broad dark dorsal abdominal cross-bands. Mostly on the apical parts of
leaves, on both upper and undersides, of unidentified Salix sp(p). in southern
California and Mexico (Hille Ris Lambers, 1966d).
 Chaitophorus 607

Chaitophorus euphraticus Hodjat Apterae pale green; BL 1.1-2.1mm.

Alatae without dorsal cross-bands. On undersides of leaves of Populus
euphraticus, attended by ants, in Iran, Jordan and Iraq; occasionally found
on P. x euroamericana and P. nigra var. italica. This aphid possibly over-
winters as parthenogenetic forms in southern Iran, possibly surviving in
December-January in fallen leaves stuck together by lepidopterous larvae
(Hodjat, 1981). Sexuales (an ovipara and an alate male) were collected on
P. x euroamericana in November in northern Iran. 2n = 22.

Chaitophorus furcatus Quednau Apterae bright or dull yellow; BL 1.2-

1.6mm. Alatae without distinct dorsal abdominal markings. In ant-attended
colonies on Salix caprea in Finland, Germany, Czechoslovakia and European
Russia. Apterous males and oviparae were collected in Finland in late
September (Pintera, 1987). 2n = 16*.

Chaitophorus gomesi Ilharco Apterae pale green mottled with dark green;
BL 1.1-1.4 mm. Alatae are undescribed. In large colonies on undersides
of leaves of Salix vitellina, attended by ants. Only known from Portugal.
Apterous males and oviparae were collected in November (Ilharco, 1968).

Chaitophorus hillerislambersi Pintera Colour of apterae in life unknown,

probably pale with dark legs; BL 2.0-2.4 mm. Other morphs are unknown.
On Populus sp. in Beijing, China (Pintera, 1987).

Chaitophorus himalayensis (Das) Apterae according to the original descrip-

tion (Das, 1918) are light green to yellowish, distinctively marked with a dark
green patch on mesothorax and two dark green crescentic bands leaving paler
areas around bases of siphunculi, meeting two elongate spots on ABD TERG
1; BL 1.1-1.9 mm. The siphunculi are short and brown. Alatae have similar
dark green dorsal markings. On Salix spp., including S. tetrasperma, feeding
along mid-ribs or leaves or on young growth, attended by ants. In northern
India, Pakistan, and possibly Thailand and China; Hille Ris Lambers (1966c)
pointed out that several records of this species from Thailand and China do
not conform to the original description. Some of these authors may have
had C. pakistanicus (q.v.). Sexuales are unknown. 2n = 18 (J. Dutta and
D.C. Gautam, pers. comm.).

Chaitophorus hokkaidensis Higuchi Colour of apterae in life unknown,

body elongate oval and probably rather dark with legs paler; BL 1.4-1.8 mm.
Alatae have broad dorsal abdominal cross-bands. Described from an uniden-
tified Salix species in Hokkaido, Japan (Higuchi, 1972).

Chaitophorus horii Takahashi Apterae whitish to pale yellow, elongate

oval; BL 1.0-1.8 mm. Alatae with black dorsal abdominal cross-bands. On
Salix, especially narrow-leafed species, in Japan, Thailand and Siberia
(Pintera, 1987; Pashchenko, 1988b). In Europe, narrow-leafed Salix have very
similar aphids, often regarded as a subspecies Ch. horii beuthani (Börner),
608 Chaitophorus

although Higuchi (1972) may have been correct in synonymizing it with horii,
as the differences in form of the dorsal hairs, etc. could be ascribed to
geographical variation. Apterous males and oviparae of the European form
occur in October. Colonies are not ant-attended (see also Heie, 1982).
Chaitophorus hypogaeus Hille Ris Lambers Apterae broadly oval, dark
dirty green, with paler greyish central area and pale antennae and legs; BL
1.4-2.0 mm. Alatae with narrow, usually interrupted, dusky to dark dorsal
abdominal cross-bands. In often large, ant-attended colonies on the subterra-
nean parts of one-year-old branches of Salix repens (= S. rosmarinifolia),
apparently restricted to plants growing in dry sandy soil (Hille Ris Lambers,
1947b; Pintera, 1987). Recorded from Belgium, The Netherlands and Poland.
Apterous males and oviparae occur in October.

Chaitophorus indicus Ghosh, Ghosh and Raychaudhuri Apterae elongate

oval, pale green; BL 1.3-2.0 mm. Alatae have dark dorsal abdominal cross-
bands. In colonies on undersides of both young and older leaves of an uniden-
tified Populus sp. (original record from Litsea is presumed to be an error).
In India (Himachal Pradesh, West Bengal). Oviparae and alate males occur
in late October (S.K. Das et al., 1981). Pintera (1987) regards Ch. manaliensis
Chakrabarti, 1977, also described from an undentified Populus sp., but with
oviparae described from P. alba (Agarwala and Mahapatra, 1990), as a possi-
ble synonym. (The chromosome number of 2n = 18 for an aphid from Salix
sp., identified as Ch. manaliensis (N.B. Pal and Khuda-Bukhsh, 1983) pre-
sumably applies to some other Chaitophorus sp. However, J. Dutta and
D.C. Gautam (pers. comm.) recorded 2n = 18 for a population from Populus
ciliata in Himachal Pradesh which agrees well with the description of Ch.
manaliensis.)
Chaitophorus inouye Pintera Apterae are pale yellow; BL 1.4-1.9 mm.
Alatae have broad black dorsal abdominal cross-bands. On leaves of Populus
spp. (koreana, maximowiczii) in Japan and also recorded from P. davidiana
and P. suaveolens in east Siberia (Kamchatka, Sakhalin, Primorskiya Kray;
Pashchenko, 1988b). Higuchi's (1972) description of Ch. dorocolus applies to
this species. Oviparae were collected on the trunk of P. maximowiczii in
October (Pintera, 1987). Ch. xizangensis Zhang (in Zhang and Zhong, 1981b)
from China is possibly inouye. 2n = 26.

Chaitophorus israeliticus Hille Ris Lambers Apterae oval, colour in life not
known, probably pale; BL 1.0-1.2 mm. Alatae and other morphs unknown.
Originally described from an unidentified Salix sp. in Israel (see Pintera,
1987); also collected from Salix alba in Turkey (BMNH colln).

Chaitophorus kapuri Hille Ris Lambers Apterae elongate oval, presumably

blackish with pale legs and a paler spinal stripe from metanotum to about
ABD TERG 3; BL 1.4-2.0 mm. Alatae have broad dark dorsal abdominal
cross-bands and wing veins conspicuously bordered with fuscous. On upper
and undersides of leaves of Populus ciliata in Murree Hills of Pakistan (Hille
 Chaitophorus 609

Ris Lambers, 1966c). Oviparae and both apterous and alate males are recorded
by Saha and Chakrabarti (1988b), who also record predators and attendant
ants.

Chaitophorus lapponum Ossiannilsson Apterae broadly oval, probably

blackish with legs paler; BL 1.7-2.3 mm. Alatae have narrow dorsal abdomi-
nal cross-bands. On leaves, petioles and shoot apices of Salix spp. in Finland
and Sweden; records from Salix purpurea in Austria and Germany refer to
Ch. mordvilkoi (see Heie, 1982; Pintera, 1987). Sexuales undescribed.

Chaitophorus leucomelas Koch (= versicolor Koch), Plate lla, b Apterae

rather elongate oval, pale green or yellow, typically with black or dark green
longitudinal pleural stripes which may be segmentally divided and dark
siphunculi; BL 1.2-2.4 mm. Alatae have dark brown dorsal abdominal cross-
bands and separate marginal sclerites. On young shoots of Populus spp. in
spring and later under leaves, in leaves stuck together by moth larvae, or leaf
galls vacated by other insects; always ant-attended. Host plants in Europe are
mainly P. nigra and related species and hybrids, but in North America a wider
range of species is colonized. Development at different temperatures was
studied by Hintze-Podufal and Thorns (1979). Oviparae and alate males occur
in October-November; sexual morph production was studied by Thorns and
Hintze-Podufal (1979). Common and widely distributed in Europe, southwest
and Central Asia; introduced into South Africa (Transvaal; BMNH colln, leg.
I.M. Millar), North and South America. 2n = 36 (Israel) or 40 (UK, South
Africa); the different numbers may indicate sibling species.

Chaitophorus longipes Tissot Apterae mainly brown, darker on head and

thorax than abdomen (Tissot, 1932b); BL 1.7-2.1 mm. Alatae have pale brown
abdomen with dark brown dorsal cross-bands and marginal sclerites. On
young tender stems of Salix caroliniana in eastern USA.

Chaitophorus longisetosus Szelegiewicz Apterae elongate oval, blackish-

brown including siphunculi, with antennae and legs also mainly dark; BL
1.4-1.9 mm. Alatae have not been found. In small colonies mainly on
older trees of Populus alba, on undersides of leaves, especially those spun
together by other insects. Oviparae, which are larger (BL 1.8-2.5 mm), occur
in the colonies in October, but males are unrecorded. In Eastern Europe
(Szelegiewicz, 1961; Pintera, 1987).

Chaitophorus longiunguis Robinson Apterae green; BL 1.6-1.9 mm. Alatae

undescribed and biology unknown. On an unidentified Salix sp. in Utah, USA
(Robinson, 1974).

Chaitophorus macrostachyae (Essig) Apterae broadly oval, greyish to

brownish or black, often with a pale central dorsal area; BL 2.0-2.8 mm.
Alatae have dark dorsal abdominal cross-bands and marginal sclerites.
Usually found on roots and parts of trunk of Salix spp. at or below ground
610 Chaitophorus

level, but colonies may also form at tips of twigs (Essig, 1912a). Four other
species have been described from this microhabitat and resembling Ch. macro-
stachyae in most characters with the exception of the degree of sclerotization
and pigmentation of the dorsum, some differences in shape and length of
hairs, and in chaetotaxy of ABD TERG 8; they are Ch. abditus (Hottes),
Ch. canens Richards, Ch. knowltoni Hille Ris Lambers and Ch. salicicorticis
(Essig). None of these differences give reliable separation of paratype and
other material in the BMNH collection, so they are all treated here as
synonyms of macrostachyae. Chaitophorus utahensis, however, synonymized
with macrostachyae by Richards (1972a), is a distinct species (q.v.).

Chaitophorus matsumurai Hille Ris Lambers (= salicicola Matsumura, 1917

nec. Essig, 1911) Apterae rather elongate oval, fuscous (according to original
description, Matsumura, 1917); BL 1.0-1.4 mm. Alatae have greenish-yellow
abdomen with dark dorsal cross-bands. Other morphs and biology unknown.
Originally described from Salix bakko in Japan and specimens from Salix
sp. in China and Korea are also this species, but many of the other host records
(as salicicola Matsumura) in the literature are probably based on misidenti-
fications (Pintera, 1987).

Chaitophorus melanosiphon Pintera Apterae short-bodied, oval, colour

in life unknown, probably pale; BL 0.8-1.7 mm. Alatae not recorded. On
Populus alba in Rumania and Iran. Oviparae and both apterous and alate
males occur in October (Pintera, 1987).

Chaitophorus minutus (Tissot) Apterae uniformly yellow according to

Tissot (1932b); BL 0.7-1.1 mm. Alatae without distinct dorsal cross-bands. In
dense colonies on undersides of leaves of Salix caroliniana in eastern USA
(Florida, Louisiana, South Carolina).

Chaitophorus miyazakii Pintera Apterae elongate oval, colour in life

unknown, probably darkish; BL 1.5-1.8 mm. Other morphs and biology not
known. On an unidentified Populus sp. in Hokkaido, Japan (Pintera, 1987).

Chaitophorus monelli (Essig) Apterae 'transparently light green throughout'

(Essig, 1912a); BL 1.0-1.2 mm. Alatae have median dorsal black patches on
pale green abdomen, only extending laterally as cross-bands on posterior
tergites. Found on tender shoots and suckers of Salix lasiolepis and S.
laevigata in California, USA. Hille Ris Lambers (1960a) redescribed this
species. Sexuales and biology are unknown.

Chaitophorus mordvilkoi Mamontova Apterae broadly oval, dark with a

yellowish-green spinal stripe from back of head to middle of abdomen; BL
1.2-2.1 mm. Alatae have dark dorsal abdominal cross-bands. In small colonies
on Salix purpurea, on shoot terminals and undersides of leaves, without ants
(Pintera, 1987). In Central, southern and Eastern Europe and eastward to
Mongolia (BMNH colln; leg. H. Szelegiewicz).
 Chaitophorus 611

Chaitophorus nassonowi Mordvilko Apterae brownish-red to brown, oval;

BL 1.5-2.8 mm. Alatae have dark dorsal abdominal cross-bands and marginal
sclerites. In ant-attended colonies on bark of young and older twigs or leaf
petioles of Populus nigra and its varieties and hybrids, in Eastern and Central
Europe. Apterous males and oviparae occur in October-November (Pintera,
1987).

Chaitophorus neglectus Hottes and Frison Apterae usually yellowish-green

to green with brownish head, prothorax and posterior abdomen and typically
with a pair of dark pleural stripes of varying width running from thorax to
ABD TERG 6; BL 1.7-2.5 mm. Alatae have dark dorsal abdominal cross-
bands and dark siphunculi. On upper and undersides of leaves, or in leaves
folded and stuck together by other insects, of aspens (Populus tremuloides,
P. grandidentata) and of P. deltoides. Colonies are often quite large, not
attended by ants (but sometimes mixed with Ch. stevensis which is ant-
attended). Oviparae and both apterous and alate males occur in September-
November. Widely distributed in North America. This aphid is very similar
to, and often confused with, Ch. populifolii, but is probably a good species;
see Richards (1972a) and Hille Ris Lambers (1960a; as populifolii sspp.
neglectus and simpsoni). 2n = 12.

Chaitophorus niger Mordvilko Apterae blackish-brown with mainly pale

antennae and legs, a thin pale ring around the base of each siphunculus
and a pale CAUDA; BL 1.2-2.3 mm. Alatae have brown dorsal abdominal
cross-bands. Living separately or in small colonies on leaves of various
Salix spp., only rarely visited by ants. Oviparae and males occur in September-
November; whether males are apterous or alate seems to depend on geo-
graphical location (Lampel, 1983). In Europe and across Asia to Siberia
(Pashchenko, 1988b), but apparently not in China or Japan. Closely related
to, and often treated as a subspecies of, Ch. salijaponicus. 2n = 30.

Chaitophorus nigrae Oestlund (= salicicola Essig) Apterae are usually

black, often with a paler spinal area which rarely extends to ABD TERG 7,
with femora of mid- and hind legs usually dark brown, but paler specimens
may also occur with variably developed dark pigmentation of metathorax
and abdomen confined to pleural areas only; BL 1.7-2.5 mm. Alatae have
dark dorsal abdominal cross-bands, often coalescent beween ABD TERG
2-6. First instar are usually black, brown or dark green (cf. Ch. pallipes).
Sometimes in large colonies on leaves and petioles of new growth of various
Salix spp. Ant attendance is not recorded. Large, pale oviparae and alate
males occur in September-November (BMNH colln). Widely distributed in
North America. This is an extremely variable species, or possibly a complex
of species, varying not only in pigmentation but in the reticulation of the
dorsal cuticle, the form of the dorsal hairs and the shape of the CAUDA.
We follow Richards (1972a) in treating Ch. salicicola and its subspecies,
described by Hille Ris Lambers (1960a), as synonyms. 2n = 24*.
612 Chaitophorus

Chaitophorus nigricantis Pintera Apterae elongate oval, colour in life

unknown but probably pale; BL 1.4-2.3 mm. Alatae have brownish dorsal
abdominal cross-bands, coalescent on ABD TERG 3-6. Biology and sexual
morphs unknown. Recorded from Salix spp. (nigricans, phylicifolia, triandra)
in Sweden and Finland (Pintera, 1987). A form on Salix sp. in Mongolia is
distinguished as a subspecies, mongolicus Pintera. Sometimes confused in the
literature with Ch. truncatus (q.v).

Chaitophorus nigricentrus Richards Apterae broadly oval, dark yellow-

brown to blackish-brown, with central part of dorsum darker than margins;
BL 1.5-1.8 mm. Alatae have dark dorsal abdominal cross-bands. Found only
on second-year growth of (unidentified) Salix sp(p).; recorded from Ontario,
Canada (Richards, 1972a) and Pennsylvania, USA (BMNH colln). Biology
and sexual morphs undescribed.

Chaitophorus nigritus Hille Ris Lambers Apterae rather elongate oval,

black with pale antennae and legs; BL 1.0-1.6 mm. Alatae not recorded. In
compact colonies around galls on the leaves of unidentified Salix sp(p). From
Pakistan and Iran (Hille Ris Lambers, 1966c; Pintera, 1987), and since iden-
tified from Salix alba in Iraq and Himachal Pradesh, India (BMNH colln).
Oviparae and one alate male were collected in Iran in December (BMNH colln;
leg. S. Hodjat).

Chaitophorus nodulosus Richards Colour of apterae in life not recorded,

probably dark brown to black; BL about 1.5-1.7 mm. Alatae and sexuales
unknown. Known only from the type series, collected on Populus sp. in Utah,
USA (Richards, 1972a).

Chaitophorus nudus Richards Apterae broadly oval, dark brown to shiny

black, with antennae and legs also dark; BL 2.0-2.6 mm. Alatae have dorsal
abdomen blackish or dark green with black spots or incomplete cross-bands,
and forewing veins black and strongly bordered, with a very conspicuous black
pterostigmal spot. In rather small colonies girdling trunk of saplings of
Populus tremuloides (Richards, 1972a). Large pale oviparae were collected in
September in Colorado (BMNH colln). Recorded from Canada (Manitoba,
Ontario) and USA (Colorado, Pennsylvania; BMNH colln).

Chaitophorus pakistanicus Hille Ris Lambers Apterae are oval, pale green;
BL 1.2-1.7 mm. Alatae are undescribed; two specimens in the BMNH collec-
tion have no dorsal abdominal markings. In dense colonies along the veins
on both sides of leaves of Salix spp. in Central and East Asia (Afghanistan,
Iran, India, Pakistan, Thailand). Oviparae were collected on S. acmophylla
in December (Hille Ris Lambers, 1966c; Pintera, 1987). Bhagat (1982b)
described a new species of parasitoid from Ch. pakistanicus in Kashmir.

Chaitophorus pallipes Richards (= albicentrus Richards) Apterae yellow

with broad black pleural stripes and femora usually pale, at least on fore and
 Chaitophorus 613

middle legs; BL 1.8-2.4 mm. Alatae have black dorsal abdominal cross-bands.
First instars are yellow (cf. nigrae). On young growth of Salix discolor (BMNH
colln; RLB) and possibly other Salix spp. in Ontario, Canada; also recorded
from British Columbia, Prince Edward Island and North Carolina (Richards,
1972a). Ant-attended. The synonymy of pallipes with saliciniger by Eastop
and Hille Ris Lambers (1976) was in error.

Chaitophorus parvus Hille Ris Lambers Apterae are black with pale anten-
nae and legs; BL 1.2-1.8 mm. Alatae with narrow dorsal abdominal cross-
bands. On undersides of leaves of Salix repens (= rosmarinifolia), attended
by ants. Recorded from Poland, Denmark, Norway and Sweden (see Heie,
1982). Pintera (1987) synonymized this species with niger, but it appears
distinct.

Chaitophorus pentandrinus Ossiannilsson Apterae are oval, pale green or

yellow; BL 1.4-2.3 mm. Alatae have dark dorsal abdominal cross-bands
and marginal sclerites. Only known from Salix pentandra in Sweden. Ant-
attended. Males are apterous (see Heie, 1982; Pintera, 1987).

Chaitophorus pheleodendroni Juchnevitch Apterae elongate oval, colour-

less; BL 1.2-1.4 mm. Alatae with brown dorsal abdominal cross-bands. In
colonies on undersides of leaves of Phellodendron amurense (Rutaceae),
introduced as an ornamental into eastern Kazakhstan (Juchnevitch, 1970).
(Possibly these were aberrant colonies of Ch. horii.)

Chaitophorus populeti (Panzer) Apterae oval, shiny dark green to black,

with antennae and legs mid-green to black; BL 1.5-2.9 mm. Alatae have broad
brown dorsal abdominal cross-bands. On young shoots and terminal leaf
petioles of various Populus spp. (especially of the alba, tremula groups)
throughout the palaearctic region. Oviparae and both apterous and alate males
occur in October-November. A subspecies, sensoriatus Mimeur, occurs on P.
alba in the Mediterranean region and is also recorded from Afghanistan (see
Pintera, 1987). 2n = 12* (RLB; Iran, China); but Pal and Khuda-Bukhsh
(1983) record 2n = 10 from India.

Chaitophorus populialbae (Boyer de Fonscolombe) Apterae short-bodied,

oval, greenish to yellowish-white, often with small green spots; BL 1.0-
2.3 mm. Abdomen of alatae with dark brown cross-bands, often coalescent
on ABD TERG 2-6. In small colonies usually on undersides of leaves of
various Populus spp. sometimes ant-attended. Throughout the palaearctic
region, in North, West and southern Africa, and introduced and widespread
in North America. Oviparae and both apterous and alate males occur in
September-November. A subspecies, yomefuri Shinji, occurs on P. alba
and P. sieboldii in Mongolia, Korea and Japan (see Pintera, 1987). Ch.
diversifolia Juchnevitch, described from P. diversifolia in Kazakhstan, is
probably a synonym of Ch. populialbae. 2n = 28 (Chen and Zhang, 1985b)
or 30.
614 Chaitophorus

Chaitophorus populicola Thomas Apterae yellow-brown to shiny black with

dark antennae and legs; BL 2.0-2.5 mm. Alatae have dorsal abdominal cross-
bars or bands and forewing veins dark and conspicuously brown-bordered.
Often in dense colonies on young shoots and developing leaves of numerous
Populus spp. throughout North America, sometimes attaining pest status in
plantations of P. deltoides (Coleman and Jones, 1988). Large, pale oviparae
and alate males occur in September-November. This species is very similar to
Ch.. nudus, but differs in chaetotaxy and feeding habit (Richards, 1972a).
A subspecies, patchae Hille Ris Lambers, was erected for populations with
long, pointed dorsal hairs. 2n = 18*, 28* and 32* in different populations
(R.L. Blackman, unpublished data), indicating that a complex of species is
involved under the name populicola.

Chaitophorus populifolii Essig (= balsamiferinus Hille Ris Lambers; =

populellus Gillette and Palmer; = essigi Gillette and Palmer) Apterae are
yellowish, sometimes with variably developed brown to black pigmentation of
head, pro- and mesothorax and ABD TERG 7 and 8, and occasionally with
a pair of dark, usually incomplete, pleural stripes; BL 1.0-2.3 mm (dwarf
apterae occur in summer and have dark legs and 5-segmented antennae).
Alatae have variably developed dark dorsal cross-bands and pale siphunculi.
On leaves of Populus spp., especially angustifolia and balsamifera, some-
times on deltoides or tremuloides (Richards, 1972a). Widespread in USA and
Canada, Mexico, and introduced into Europe (Germany), probably on balsam
poplar cuttings (Pintera, 1987). Oviparae and both apterous and alate males
occur in September-November. There has been considerable confusion of
nomenclature of this taxon - see Sorensen (1989). 2n = 12.

Chaitophorus pruinosae Narzikulov Apterae are dark brownish-green; BL

1.6-2.1 mm. On undersides of leaves of Populus pruinosa in Tadzikhistan and
also collected from P. nigra in Afghanistan (BMNH colln). Males are alate.
See Pintera (1987).

Chaitophorus pusillus Hottes and Frison Apterae whitish, BL 0.9-1.3 mm.

Alatae with dark dorsal cross-bands coalescing into a central patch. On upper
and undersides of leaves of Salix amygdaloides in Illinois, USA (Hottes and
Frison, 1931), and also recorded from 5. discolor and unidentified Salix spp.
in other parts of USA and Canada.

Chaitophorus pustulatus Hille Ris Lambers Appearance in life not recorded

but apterae probably variably pigmented, brown to black with paler central
dorsal area; BL 1.0-1.9 mm. Alatae not described; those in the BMNH
collection have dark dorsal abdominal cross-bands. On leaves of Salix sp.
in Utah, Coloradoa and Wyoming (Hille Ris Lambers, 1960a), and also
collected in British Columbia (Richards, 1972a), Idaho and Oregon (BMNH
colln) - in all cases from unidentified Salix. Small pale oviparae and apterous
males have been collected in Colorado in October (BMNH colln, leg. P.C.
Hottes).
 Chaitophorus 615

Chaitophorus quinquemaculatus Bozhko Apterae are elongate oval, whitish

with three longitudinal stripes and two (?pleural) spots (Bozhko, 1976); BL
1.2-2.2 mm. Alatae have brownish dorsal abdominal cross-bands. In dense
colonies on twigs, or rarely on undersides of leaves, of Salix acutifolia and
also recorded from S. caprea and S. fragilis. Pintera (1987) regards it as a
subspecies of Ch. saliapterus.

Chaitophorus ramicola (Börner) Apterae are broadly oval, mainly very dull
greyish-olive to black with a more-or-less distinct paler spinal stripe and dark
antennae and legs; BL 1.4-2.6 mm. Alatae have narrow dorsal abdominal
cross-bands. In ant-attended colonies on bark of twigs of mainly broad-leaved
Salix spp., in northern and Central Europe. Apterous males and oviparae
occur in October (Heie, 1982). Chaitophorus brunealineatus Juchnevitch,
1970, described from S. russica in Kazakhstan, is probably a synonym.

Chaitophorus remaudierei Pintera Apterae are elongate oval, BL 1.4-

2.0mm, probably variably dark-pigmented but colour in life not recorded.
Alatae are undescribed. On an unidentified Salix sp. in Iran (Pintera, 1987).
Apterous males and oviparae were collected in October.

Chaitophorus saliapterus Shinji Apterae are elongate oval, pale green; BL

1.0-2.0 mm. Alatae have dark dorsal abdominal cross-bands. On undersides
of leaves of Salix spp. (especially integra), attended by ants, in Siberia
(Pashchenko, 1988b), Japan and Korea (Pintera, 1987). 2n = 30 (Shinji's 1931
record of 2n = 14 (n = 7) should probably be applied to another species of
Chaitophorus).

Chaitophorus saliciniger (Knowlton) Apterae variably pigmented but often

black, with a paler, spindle-shaped, spinal stripe; BL 1.7-2.2 mm. Alatae have
dark dorsal abdominal cross-bands. On bark of twigs of unidentified Salix
sp(p). in Colorado and Utah, USA.

Chaitophorus salicti (Schrank) Apterae black with a pale spinal stripe in

spring, but small apterae in late summer are yellowish-white with reddish-
brown or greyish-black dorsal markings; BL 1.3-1.8 mm. Alatae dark, with
broad dorsal abdominal cross-bands. On undersides of leaves of broad-leafed
Salix spp., usually along main veins, attended by ants. Throughout Europe,
east to Iran and also recorded from China (Zhang et al., 1986a). Apterous
males and oviparae occur in September-October (Heie, 1982; Pintera, 1987).
An internal cecidomyid parasite (?Endaphis sp.) was observed emerging from
the anus of an aphid of this species in June 1952 at Gulpen, The Netherlands
(Nijveldt, 1969). 2n = 28.

Chaitophorus salijaponicus Essig and Kuwana Apterae are presumably dark

brown or black, sometimes with paler narrow spinal stripe; BL 1.3-1.9 mm.
Alatae have dark dorsal abdominal cross-bands. On leaves of Salix spp. in
Japan, China and Siberia. Very similar to Ch. niger, which is often treated
616 Chaitophorus

as a subspecies. Two other forms in this group are distinguished as sub-

species by Pintera (1987); ssp. stroyani from S. phylicifolia in Italy, and ssp.
szelegiewiczi from Salix sp. in Mongolia.

Chaitophorus saliniger Shinji Apterae shining blackish-brown to black with

legs and distal parts of antennae dark and CAUDA conspicuously paler; BL
1.0-1.8 mm. Alatae have broad black dorsal abdominal cross-bands. On
young leaves, petioles or shoot tips of narrow-leafed Salix spp., often ant-
attended, in Japan, China and Korea. Apterous males and oviparae occur in
November-December (Higuchi, 1972; Pintera, 1987). 2n = 8.

Chaitophorus shaposhnikovi Mamontova Apterae whitish with five greenish

spots on dorsum (Mamontova, 1955); BL c. 1.2-1.3 mm. Alatae undescribed.
On undersides of leaves, along main veins, of an unidentified Salix sp. in
Ukraine and west Kazakhstan (Shaposhnikov, 1964a).

Chaitophorus similis Pintera Colour of aptera in life unknown, presumably

pale; BL 0.8-1.3 mm. Alatae unknown. On Salix incana in southern France
(Pintera, 1987). Apterous males in September.

Chaitophorus stevensis Sanborn (= delicatus Patch) Apterae are pale green

with darker dorsal markings; BL 1.6-2.0 mm. Alatae have black head and
thorax, greenish abdomen with dark cross-bands often coalesced into a central
patch and darkish siphunculi (Richards, 1972a). Ant-attended, on leaves of
Populus spp., apparently restricting its feeding to aspens and cottonwoods;
records from balsam poplars are probably misidentified populifolii. It is
frequently associated with leaf-fold galls of Thecabius populiconduplifolius.
Widely distributed in North America. 2n = 14.

Chaitophorus tremulae Koch Apterae are elongate oval, dark brown to

black, sometimes with a paler spinal stripe, legs brown with hind pair darker;
BL 1.2-2.5. Alatae have very broad and nearly coalescent dorsal abdominal
cross-bands. Immatures are bright green. In small colonies on undersides of
leaves of P. tremula and a few related species, often in leaves spun together
by other insects, or in leaf-nest galls of Pemphiginae. Not usually ant-
attended. Oviparae and alate males occur in October. Gärdenfors (1986)
described a specific parasitoid in Sweden. Chaitophorus tremulae occurs
throughout Europe (Heie, 1982; Pintera, 1987). Pintera classified two related
East Asian forms as subspecies of tremulae; shantungensis Tseng and Tao,
1936 on Populus spp. in China and Japan, and a pale form living on leaves
or in Pemphigus galls on Populus laurifolia in Japan and Mongolia (ssp. sorini
Pintera). 2n = 18*.

Chaitophorus truncatus (Hausmann) Apterae are elongate oval, pale green

in spring and often with three dark green longitudinal stripes, but in summer
sometimes much darker or even blackish, with mainly pale legs and antennae;
BL 1.2-2.4 mm. Alatae with variably-developed dorsal abdominal markings,
 Chaitophorus 617

usually not forming solid cross-bands. In small colonies on leaves of various

narrow-leafed Salix spp., throughout Europe and east to Iran and western
Kazakhstan (Heie, 1982; Pintera, 1987). Not ant-attended. Apterous males
and oviparae in October-November. 2n = 30.

Chaitophorus utahensis (Knowlton) Apterae broadly oval, colour in life not

noted but presumably rather dark; BL 2.0-2.5 mm. Alatae have dark dorsal
abdominal cross-bands and dark wing veins with narrow fuscous borders. On
bark of twigs and extending onto leaf petioles, or on trunk 15-20 cm above
ground level (BMNH colln; leg. G.F. Knowlton), of unidentified Salix sp(p).,
ant-attended, in western USA (Utah, Colorado, California, Idaho, Oregon;
Hille Ris Lambers, 1960a and BMNH colln).

Chaitophorus variegatus Szelegiewicz Apterae greenish-yellow with some-

times darker head and thorax; BL 1.4-2.3 mm. Other morphs unknown.
On undersides of leaves of Populus koreana in Korea, attended by ants
(Szelegiewicz, 1981).

Chaitophorus viminalis Monell Apterae of varying pigmentation in life,

usually pale green to yellow in spring but often dark brown to black in
summer; BL 1.3-1.9 mm. Alatae have dark dorsal abdominal cross-bands
often coalesced into a central patch. On leaves and young growth of various
narrow-leafed Salix spp. throughout North America. Oviparae and alate
males occur in October-November. Probably often misidentified in the
literature; differences from similar-looking species (e.g. nigrae, pusillus)
are discussed by Hille Ris Lambers (1960a). See also Richards (1972a).
2n = 18.

Chaitophorus viminicola Hille Ris Lambers Apterae rather elongate oval,

brown or blackish, usually with a paler spinal stripe from pronotum to about
ABD TERG 3; BL 1.9-2.5 mm. Alatae have broad dark dorsal abdominal
cross-bands. On Salix nigra (BMNH colln) and unidentified Salix sp(p). in
eastern USA and Canada (Ontario). Biology and sexuales unknown.

Chaitophorus vitellinae (Schrank) Apterae broadly oval, yellowish-green

with two broad darker green to greenish-brown pleural longitudinal stripes,
and pale antennae and legs; BL 1.3-2.3 mm. Alatae have dark dorsal abdom-
inal cross-bands. On bark of young twigs and leaf petioles of narrow-
leafed Salix spp., especially S. alba; attended by the ant Lasius fuliginosus.
Throughout Europe except Iberian peninsula. Pintera (1987) described a
subspecies (danubicus), with a distinctly knobbed CAUDA and thickened dor-
sal hairs, from S. alba in Rumania, Czechoslovakia and Turkey. Apterous
males and oviparae occur in September-October (Heie, 1982).
618 Chaitoregma

CHAITOREGMA Hille Ris Lambers and Basu

Hormaphidinae: Cerataphidini

Bamboo-feeding aphids characterized by the rounded frontal horns and the

heavily sclerotized dorsum.

Chaitoregma tattakana (Takahashi) Apterae yellowish-brown to deep violet-

black, covered with fine wax dust; BL 1.5-1.9 mm. In large colonies on under-
sides of leaves of Bambusa and Arundinaria spp. Recorded from India and
Taiwan (Hille Ris Lambers and Basu, 1966; Liao, 1976). Chaitoregma
aderuensis (Takahashi, 1935f), from Bambusa sp. in Taiwan, is not clearly
distinct. Life cycle is undescribed, but galls of a Chaitoregma sp. occur on
Ligustrum in Hong Kong (BMNH colln, leg. C.S.K. Lau).

CHROMAPHIS Walker Drepanosiphinae: Phyllaphidini

Three species of small pale yellow aphids, related to Panaphis and asso-
ciated with Juglans. Quednau (1973) and Chakrabarti (1988) revised the two
palaearctic species, which are also both recorded from Aleurites moluccana
(Euphorbiaceae), presumably acting as a 'reserve host'. Trioxys pallidus
and Praon flavinode specialize on Chromaphis and the related genera
Eucallipterus, Myzocallis, Tinocallis and Tuberculatus (Stary, 1976). Aphe-
linus perpallidus attacks Melanocallis and Monellia as well as Chromaphis.

Chromaphis californica (Essig) Adult viviparae all alate, yellow with dark
antennae and tibiae, and with a short narrow black marginal stripe on each
side of prothorax; BL 1.7-2.0 mm. Immatures and oviparae (collected in
November) have banded antennae. Main host is Juglans californica in
California (Richards, 1968b) and Utah (BMNH colln); there are also records
(perhaps 'casual' occurrences) from Carya illinoensis and Pterocarya spp.
(Walker et al., 1978).

Chromaphis hirsutustibus Kumar and Lavigne Adult viviparae all alate,

pale yellow, often with black patches on ABD TERG 4 and 5, and with dark
spots at ends of wing veins; BL 1.6-12.5 mm. On Juglans regia in India, Nepal
and China (Zhang and Zhong, 1985c); both immatures and adults are recorded
also from Aleurites moluccana in India and China. Chakrabarti (1988)
described the oviparae, collected in September. The sexual morphs described
under this name by Chakrabarti (1978) were C. juglandicola.
 Chuansicallis 619

Chromaphis juglandicola (Kaltenbach) Adult viviparae all alate, pale lemon

yellow or yellowish-brown with pale brown thoracic lobes and (in autumn)
paired brown spots on ABD TERG 4 and 5; BL 1.2-2.3 mm. Living scattered
under leaves on Juglans regia, with occasional records from other Juglans
spp., and there is also a record from Aleurites moluccana (of oviparae;
Chakrabarti, 1978, as hirsutustibus). In Europe, Central Asia, India, Pakistan,
China and North America. Oviparae and males occur in May-June in India,
and August-November in Europe, but not before late September in USA.
Biology and ecology have been studied intensively, especially in California
(e.g. Nowierski and Gutierrez, 1986), where biological control has been suc-
cessful (Hoy and Cave, 1988). See also B & E, 1984, p. 259. 2n = 8.

CHROMOCALLIS Takahashi Drepanosiphinae: Phyllaphidini

One or more species on Ulmus in East Asia. They are plump-bodied aphids
with rather short antennae, thick dark hairy tibiae and a deeply cleft anal
plate.

Chromocallis nirecola (Shinji) Adult viviparae all alate, green, with black
tibiae and forewing veins bordered with fuscous; BL 3.0-3.2mm. On Ulmus
spp. (japonica, pumila) in Japan, Korea, China and Siberia (Higuchi, 1972;
Pashchenko, 1988b). Biology is little known; alate males were collected in
September in Japan (BMNH colln, leg. M. Sorin), and Paik (1972) recorded
oviparae and males in Korea. Chromocallis pumili Zhang in Zhang and
Zhong, 1982c and Ch. similinirecola Zhang in Zhang and Zhong, 1982c,
described from China, do not seem clearly distinguishable from nirecola.
2n = 18*.

CHUANSICALLIS Tao Drepanosiphinae: Phyllaphidini

A genus for one East Asian species with a well-defined median suture on the
head, a very long third antennal segment with regularly-spaced, slit-like secon-
dary rhinaria and a rather short processus terminalis.

Chuansicallis chengtuensis Tao All adult viviparae are alate, green, secret-
ing profuse cottony wax threads, with dark hind legs and wings held hori-
zontally over the abdomen when at rest; BL 1.6-2.0 mm. The life cycle and
host relationships are unclear; numerous alate males and apterous oviparae
were described from Rhus semialata ( = javanica) in December in Taiwan
(Tao, 1963), but alate viviparae have been collected from Dalbergia hupeana
620 Cinara

in November in Taiwan, and also in May-June in China (Kwangtung,

Szechuan; Zhang and Zhong, 1980c, as Sinotherioaphis pterothorax). Alatae
were also recorded from Phyllanthus sp. in Bhutan (A.K. Ghosh and
Quednau, 1990).

CHUCALLIS Tao - Drepanosiphinae: Phyllaphidini

A genus for one East Asian bamboo-feeding species with very long dorsal
abdominal tubercles.

Chucallis bambusicola (Takahashi) Adult viviparae all alate, dark purple to

black with pale antennae and legs and clear wings; BL 1.6-2.0 mm. On shoots
and undersides of leaves of bamboos (Bambusa stenostachya, Dendrocalamus
latiflorus) in Taiwan and China (Kwangtung, Szechuan; Tao, 1964). Very
active, jumping when disturbed. Anholocyclic in Taiwan (Takahashi, 1923; as
Myzocallis bambusicola), and sexual morphs are unrecorded.

CINARA Curtis Lachninae: Cinarini

A very large genus resulting from a highly successful adaptive radiation on

Coniferae of the families Pinaceae and Cupressaceae. As understood here,
Cinara includes Cedrobium and Cupressobium as subgenera and contains
about 200 described species, about 150 of them native to North America, 30
in Europe and the Mediterranean, and 20 in the Far East. Many of the North
American species were described in a long series of short papers by Hottes in
1951-1964, and a number of these are only known from the original descrip-
tion. The European species fall into fairly well-defined groups that have been
dignified as separate genera or subgenera, but there are difficulties with apply-
ing most of these groupings to the world fauna, possibly due in part to the
inadequate descriptions available for many species, and they are not used here.
All Cinara live without host alternation on the roots, trunk, branches,
twigs, shoots or foliage of their hosts. They are often regularly attended by
ants and have specialized parasitoids, Pauesia spp. (Smith, 1944, as Pro-
taphidius; Starý, 1976). Males may be apterous or alate depending on species,
and oviparae often differ from viviparae in the presence of a perianal ring of
wax.
For an introduction to the genus see Eastop (1972). Regional accounts are
available for Britain (Eastop, 1972; Carter and Maslen, 1982); Central Europe
(Pintera, 1966; Szelegiewicz, 1962); Germany (P.P. Müller, 1969; Heinze,
1962); Austria (Fossel, 1970); Italy (Binazzi, 1978); Spain (Gutierrez et al.,
 Cinara 621

1985); India (A.K. Ghosh, 1982b); Siberia (Pashchenko, 1988b); Korea (Paik,
1972); Japan (Inouye, 1941, 1956, 1970); New Zealand (Baker, 1986);
eastern Canada (Bradley, 1951); eastern USA Pinus-feeders (Pepper and
Tissot, 1973); and US Rocky Mountain region (Palmer, 1952). Danielsson and
Carter (1993a) keyed the subgenus Cupressobium. Voegtlin and Bridges (1988)
provide a catalogue and bibliography of the North American species.

Cinara abietihabitans Zhang and Zhong BL of aptera c. 6.0mm; appear-

ance in life not recorded. Described from Abies sp. in Yunnan, China (Zhang
and Zhong, 1985e). Related to C. conflnis but with longer antennae, shorter
hind tibial hairs and a longer R IV.

Cinara acadiana Hottes Appearance in life unknown: BL of aptera 2.9-

3.2mm. Described from Picea glauca in New Brunswick, Canada (Hottes,
1956b), and not found since.

Cinara acutirostris Hille Ris Lambers Dark brown to bronze, with a pattern
of wax dust similar to C. pini; BL of aptera 2.6-3.6 mm. On twigs of Pinus
nigra and P. pinea in Western, southern and Central Europe (England,
France, The Netherlands, Bulgaria, Czechoslovakia, Italy, Spain, Portugal).
Alatae are produced in May-June (Italy) or June-July (England). Apterous
males and oviparae occur in October-November, and eggs are laid on the
needles (Eastop, 1972; Binazzi, 1973).

Cinara alacra Hottes and Essig Colour in life not known; BL of aptera
4.6 mm. Described from a single specimen on Abies concolor in Nevada, USA
(Hottes and Essig, 1953b). There is also a record from California (Smith and
Parron, 1978).

Cinara anzai Hottes and Essig Apterae yellowish-brown, varying in dark-

ness in different individuals, with interrupted white powdery dorsal abdominal
cross-bands giving a whitish or brownish mottled appearance. Siphuncular
cones dark each surrounded by a pale area, appendages light brownish-yellow
with distal parts of femora, tibiae and tarsi dark; BL 3.5-3.9 mm. On branches
of Pinus albicaulis in California and Oregon, USA. Oviparae and alate males
were collected in September (Hottes, 1956e).

Cinara apacheca Hottes and Butler Probably uniformly dark brown and
shining; BL of aptera about 2.9-3.0 mm (Hottes and Butler, 1955). On Pinus
edulis in Arizona, USA. Not collected since original description, biology and
sexuales unknown.

Cinara apini (Gillette and Palmer) Apterae dark brown mottled with wax
powder on spinal and marginal areas and intersegmentally, with appendages
mainly yellowish to dusky, siphunculi and distal two thirds of tibia black; BL
3.2-4.0 mm. In large colonies on bark of twigs and small branches of Pinus
flexilis (Palmer, 1952), and perhaps sometimes on P. albicaulis (one aptera
622 Cinara

in BMNH colln), in western North America. (The record of this species from
P. banksiana in Quebec (Quednau, 1966) is a misidentification of C. bank-
siana.) Oviparae and alate males occur in September-October, and eggs with
a light covering of wax-wool are laid on the needles.

Cinara arizonica (Wilson) Apterae dark brown to greyish-black with patches

of wax powder on thorax and sides of abdomen, the largest ones being just
anterior to siphunculi and tending to extend across ABD TERG 4 (Palmer,
1952; Voegtlin, 1976); BL 4.5-6.0 mm. On bark of stems of P. ponderosa,
and there are also arboretum records from several other pines in subsections
Ponderosae, Sabinianae and Australes. In western North America.

Cinara atlantica (Wilson) (= C. carolina) Apterae are greyish-brown with

dark brown head, pro- and mesothorax, and paired dark brown irregular
dorsal patches on metathorax and ABD TERG 1, extensive more-or-less
circular dark brown siphuncular sclerites, and legs dark except for a pale
section on basal part of each tibia; BL 2.2-3.3 mm (Pepper and Tissot, 1973).
Forming dense colonies in spring on new growth, or on bark of small branches
among the needle bases, of many Pinus spp., mostly in subsection Australes.
In eastern USA (especially common in southeast) and Canada (Ontario), and
in Cuba (Hernandez and Rodriguez, 1985) and Jamaica (BMNH colln). Not
hitherto recognized to occur in western USA, but the BMNH has large series
of what appears to be this species from numerous Pinus spp. in Los Angeles
Co. Arboretum at Arcadia, California (leg. H.G. Walker). Oviparae and alate
males occur in October, and eggs are laid on the needles. Anholocycly occurs
in warmer regions. Oliver and Chapin (1988) summarize biology and control
of C. atlantica in P. virginiana plantations in Louisiana. Patti and Fox (1981)
studied seasonal trends, and Carner et al. (1977) recorded fungal attack, in
populations on P. taeda in South Carolina. 2n = 10.

Cinara atra (Gillette and Palmer) Apterae are rather long-bodied with
narrow head and prothorax, shining black or dark brown without any wax;
BL 2.0-3.0 mm. Immatures are orange-yellow to olive-brown or reddish. In
small colonies on smooth bark of small branches of young pinyon pines, Pinus
edulis and P. monophylla, in Colorado and Utah, USA. Apterous males and
oviparae occur in October-November (Palmer, 1952).

Cinara atripes Hottes Colour of apterae in life not recorded, probably dark
brown or black, with wholly dark legs; BL 1.6-2.3 mm (Hottes, 1958a). On
Picea glauca in Maine, USA. Biology and other morphs unknown, and
apparently not collected since the original description.

Cinara atroalbipes David, Narayanan and Rajasingh Apterae are green to

dark green with a longitudinal spinal pale streak, appendages mainly pale
brown except for the wholly black fore tibiae; BL 2.7-3.2 mm. Originally
described from 'Pine' (David et al., 1970) in Himachal Pradesh, India, but
the host plant of apterous specimens from the type locality in the BMNH
 Cinara 623

collection (leg. A.N. Chowdhuri) was identified as Picea smithiana, and this is
probably the true host. Biology and sexual morphs unknown.

Cinara atrotibialis David and Rajasingh Apterae are brown with darker
antennae and legs, the tibiae and tarsi almost black; BL 3.3-4.3 mm. On young
shoots at needle bases of Pinus kesiya (= insularis), and also recorded from
P. roxburghii (A.K. Ghosh, 1982b). In India (Himachal Pradesh, Meghalaya),
and probably as geographic variants in Thailand (as C. khasyae Robinson) and
the Philippines (Eastop, 1976). Agarwala (1989) studied population trends
in the field, Samanta and Raychaudhuri (1984) reported the effects of a
parasitoid (Pauesia laricis) and Agarwala (1988) studied development in the
laboratory. Sexuales and life cycle are unknown; active stages are present in
December, so it is presumably at least partially anholocyclic. 2n = 10 (Kurl,
1988).

Cinara azteca Hottes Appearance in life unknown; BL of aptera 2.0-2.3 mm

(Hottes and Essig, 1954b). On Pinus ?oocarpa in Mexico. Not recorded since
the original description; possibly these were small specimens of C. atlantica.

Cinara balachowskyi Remaudière Apterae rather elongate oval with narrow

head and thorax, chocolate brown with uniformly dark legs and a light dusting
of white wax powder on the ventral abdomen (Remaudière, 1974; Melia
Masia, 1978); BL 3.3-4.9 mm. In compact colonies in bark crevices on the
trunk of Pinus halepensis in the western Mediterranean (France, Spain). Life
cycle and sexuales are unknown; active stages have been collected in Decem-
ber, suggesting anholocycly.

Cinara banksiana Pepper and Tissot Apterae have dark head and thorax
and paler brown abdomen with paired dark markings and large, prominent
shiny black siphuncular cones; BL 2.4-3.6 mm. On previous year's growth or
in dense colonies on new shoots, producing numerous alatae in spring and
early summer; on Pinus banksiana, and apparently specific to this host, in
Canada and northern USA. Oviparae and alate males occur in October and
eggs are laid on the needles. Bradley and Hinks (1968) studied the relationship
with ants in Manitoba.

Cinara bonica Hottes Colour in life not recorded, probably brownish with
some wax dust; BL 3.4-3.6 mm. Collected in Alaska on an unknown host, pro-
bably either Picea glauca or P. mariana and not found since (Hottes, 1956g).

Cinara bonita Hottes Colour in life not recorded, probably brownish with
some wax dust; BL about 5.7 mm. Collected in Alaska on an unknown host,
probably either Picea glauca or P. mariana, and not found since (Hottes,
1956g, 1964).

Cinara braggii Gillette Apterae in life covered in white wax powder, with
legs pale; BL 3.2-3.8 mm. Feeding solitarily on bark of twigs and on new
624 Cinara

growth of Picea spp., with a wide distribution in North America. Numerous

alatae are produced in the second and third generations. Oviparae and alate
males in October (Gillette, 1917). 2n = 10.

Cinara brauni Börner Apterae golden-brown with a dusting of wax powder,

and usually with an extensive dark brown to black patch on ABD TERG 5-7
encompassing the siphunculi, although this patch is reduced in some popula-
tions (Binazzi, 1978); BL 2.7-3.8 mm. On current year's growth and one-year-
old twigs of Pinus nigra, attended by ants (usually Formica rufa), in Europe
and east to Turkey and Crimea. Oviparae and alate males in October-
November. Scheurer (1971b) described the fundatrix, alata and sexuales.
Binazzi and Roversi (1987) studied seasonal differences in abdominal scleroti-
zation, including holocyclic and anholocyclic populations.

Cinara brevipilosa Voegtlin, Remaudière and Peña Martinez Colour of

apterae in life unknown, probably dark, with an extensive black dorsal
abdominal sclerite that does not however encompass the small siphuncular
cones; BL 2.0-3.5 mm. A species with very short dorsal abdominal hairs, col-
lected from an unidentified Pinus sp. in Mexico (Voegtlin et al., 1986). Biology
unknown.

Cinara brevispinosa (Gillette and Palmer) Apterae are brown lightly dusted
with wax powder medially and marginally; BL 3.0-3.5 mm. In colonies on
bark of twigs and small branches of Pinus contorta (incl. var. murrayana) in
western North America. Oviparae and alate males in late August-October
(Gillette and Palmer, 1931). Cinara sclerosa Richards is a synonym.

Cinara burrilli (Wilson) Apterae are, according to the original description,

black 'with pruinose patches that produce a calico effect' (but see Hottes,
1951); BL 2.5-3.0 mm. On bark of twigs of Juniperus spp. in the Rocky
Mountain region, USA (Palmer, 1952). Biology and sexuales unknown.
Cinara pulverulens may be a synonym.

Cinara caliente Hottes Apterae are dusky grey, variably dusted with wax
powder, with black sclerotization of thorax and ABD TERG 1 often in form
of an inverted 'V' (Hottes, 1955c); BL 1.6-2.2 mm. On bark of twigs, espe-
cially on needle-free sections, of Pinus edulis and P. monophylla in western
USA (Hottes, 1960b). Oviparae and alate males in September.

Cinara californica Hottes and Essig Appearance in life unknown, in cleared

and mounted specimens all tibiae are uniformly dark (Hottes and Essig,
1953a); BL 3.1-3.5 mm. This species is something of a mystery, being
described from the Chinese pine P. tabulaeformis in California. Long-haired
Cinara on other Pinus spp. from California in the BMNH collection, including
H.G. Walker's material previously identified as californica (Walker et al.,
1978), have two-toned tibiae and short tarsi and seem to be the East Coast
species atlantica (q.v.). Cinara californica is included in the key to species
 Cinara 625

from oriental Pinus, in case it is a previously unrecognized oriental species

introduced into California.

Cinara canatra Hottes and Bradley Shining black, without wax; BL 2.3-
3.5 mm. In spring in dense colonies on bark of smaller branches some distance
from the tips, later moving to form clusters on the bark of the trunk (Hottes
and Bradley, 1953). Probably specific to Pinus banksiana, where it may occur
in mixed colonies with, and be confused with, C. nigra (Pepper and Tissot,
1973); records from other hosts are referable to C. gracilis or C. nigra. In
Michigan, USA, and Ontario, Manitoba and Newfoundland, Canada.
Oviparae in October, males unrecorded.

Cinara caudelli (Wilson) Appearance in life unknown and not found since
original collection, on an unidentified Picea sp. in British Columbia (Wilson,
1919b). BL of aptera about 2.4mm. Life cycle and sexuales unrecorded.

Cinara cedri Mimeur Apterae dark bronze or reddish-brown with a light

dorsolateral and ventral dusting of wax powder; BL 2.5-3.8 mm. In compact
colonies on branches of Cedrus spp. in Europe (UK, France, Italy, Spain),
North Africa (Morocco), southwest Asia (Iran, Turkey) and recently found
in Argentina (BMNH colln, leg. M.A. Delfino). Oviparae and alate males
occur in October, sometimes concealed under growths of lichen on the
branches (Carter and Maslen, 1982). See Covassi and Binazzi (1974) or
Stroyan (1979) for a fuller account. 2n = 10.

Cinara cembrae (Seitner) Apterae are shiny brown, BL 3.3-4.8 mm. On

bark of two-year-old or older parts of branches of Pinus cembra in moun-
tainous areas of Central Europe (Austria, Czechoslovakia, Switzerland) and
in Russia (Leningrad district). Oviparae and both apterous and alate males are
produced in late August (Pintera, 1966). Records of cembrae from East Asia
should probably all be referred to C. mongolica.

Cinara chibi Inouye Apterae have brown head and thorax and shiny dark
brown to black abdomen, with a pale brown spinal stripe (Inouye, 1962); BL
2.4-3.1 mm. On stems of young trees, or on three- to four-year-old parts of
branches, of Larix spp. in Japan. Oviparae and alate males in October
(Inouye, 1970). Very similar to C. cuneomaculata.

Cinara chinookiana Hottes Apterae are shiny apple green to brownish-

green with brown-black head and dark markings on thorax and posterior
abdomen; BL 3.3-3.5 mm. On Abies lasiocarpa and A. amabilis (BMNH
colln), living singly on upper sides of small twigs near their tips (Bradley,
1961). In western North America. Apterous males and oviparae in October
(Hottes, 1957b).

Cinara cognita Hottes and Essig Appearance in life unknown: BL of

aptera 3.6mm (one specimen), BL of alatae 2.4-3.3 mm (Hottes, 1955a). On
626 Cinara

Juniperus communis in western USA (California, Washington). Biology and

sexuales not known.

Cinara coloradensis (Gillette) (= caliginosa Hottes) Apterae have abdomen

dullish black dorsally, somewhat shiny black ventrally, and reddish-brown
head and thorax (Gillette, 1917); BL 2.2-3.4 mm. In large, dense colonies on
undersides of small two- or three-year-old, branches and on upper trunk
(Bradley, 1961) of Picea spp. Widely distributed in western USA and across
Canada. 2n = 10.

Cinara comata Doncaster Apterae pale brown; BL 4.1-4.5 mm. On Picea

smithiana in northwest India (A.K. Ghosh, 1982b). Biology and sexual
morphs unknown.

Cinara commatula Hottes and Essig Appearance in life not recorded; BL of

aptera 4.3-4.7 mm. One aptera and one alata were described, from Pseudo-
tsuga taxifolia in California, USA (Hottes and Essig, 1954b).

Cinara confinis (Koch) (= abieticola Cholodkovsky) Apterae and alatae are

large, dark brown or greenish-black with a double row of blackish slightly
shining spots and small flecks of wax powder in transverse rows (Carter and
Maslen, 1982); BL 3.8-7.8 mm. Usually in spring colonies on trunk, branches
and twigs of Abies spp. and in summer on the roots; sometimes also on Cedrus
spp., where it may build up large spring colonies on trees growing in urban
areas. Recorded from Europe, Central Asia, India, Pakistan, Siberia (Pash-
chenko, 1988b; as piceae), Canada, western USA and Argentina (Blanchard,
1939; as grossa). Struble et al. (1976) studied its biology in Maine, USA.
Oviparae and alate males are recorded in September-October in northern
Europe, Pakistan and North America, but are not recorded from other coun-
tries where the species is common, suggesting that anholocycly occurs in mild
winter climates (see also Eastop, 1972). Heikinheimo (1963) records natural
enemies in Finland. Cinara tenuipes, described as a subspecies of confinis, is
here regarded as a distinct species. 2n = 12*.

Cinara contortae Hottes Colour in life not recorded; BL 2.5-3.2 mm. On

Pinus contorta in California (Hottes, 1958d). Biology and sexuales unknown.
This species is keyed on the basis of specimens in the BMNH collection (leg.
D.J. Voegtlin) which, unlike the original description, have well-developed
anterior dorsal abdominal sclerites and in this respect resemble C. medi-
spinosa, but have much shorter dorsal abdominal hairs. This species is also
very close to C. brevispinosa and C. banksiana; taxonomie relationships in
this group of species require investigation.

Cinara costata (Zetterstedt) Apterae are wax-covered, light brown or yellow-

brown, sometimes with a dull metallic golden sheen and with a pair of dark
bottle-green dorsal longitudinal stripes which sometimes coalesce at about the
level of the siphunculi (Carter and Maslen, 1982); BL 2.7-3.8 mm. Alatae have
 Cinara 627

characteristically pigmented forewings with the media only once-branched. In

small colonies on smaller woody twigs (which receive a deposit of mealy wax)
on lower branches of Picea spp., usually not attended by ants (Pintera, 1966;
but see Eastop, 1972). It occurs in Europe, East Asia, Australia, Greenland,
Canada and USA. Oviparae and males in October. 2n = 10.

Cinara covassii Binazzi Apterae shimmering light bronze; BL 4.1-5.0 mm.

On older branches of Pinus mugo in the eastern Alps, northern Italy. Ovipara
in August, alate viviparae and males unknown (Binazzi, 1991a).

Cinara cronartii Tissot and Pepper Apterae are dark brown with mainly
dark legs; BL 2.6-4.2 mm. This aphid seems normally to restrict its feeding
site to lesions and cankers of the rust Cronartium fusiforme on trunks and
branches of Pinus spp. (taeda, elliotti, clausa, serotina) in eastern USA (Pep-
per and Tissot, 1973). Introduced into South Africa, where severe infestations
cause economic damage to P. taeda (Raubenheimer and Shaw, 1987). Biology
was studied in the laboratory in South Africa by van Rensburg (1989), and
biocontrol with a parasitoid (Pauesia sp.) has been attempted (Kfir et al., 1985;
Kfir and Kirsten, 1991). Sexuales are unrecorded; in Florida the aphids remain
in rust lesions throughout the year. 2n = 10*.

Cinara cuneomaculata (Del Guercio) (= boerneri) Apterae and alate of

variable colour, darkish brown to orange-reddish, sometimes with dark green
segmental markings and with a ventral dusting of greyish wax powder that
sometimes extends dorsally (Carter and Maslen, 1982); BL 2.4-4.6 mm. In
small colonies on young twigs and shoots of Larix spp., usually but not always
ant-attended, in Europe and eastward to Mongolia (see Eastop, 1972; as
boerneri) and Siberia (Pashchenko, 1988b). Oviparae in October, alate males
have been recorded as early as June. It is virtually indistinguishable from
C. laricicola in Japan (Inouye, 1962), which may be a synonym. 2n = 10.

Cinara (Cupressobium) cupressi (Buckton), Plate 16b Apterae and alatae

orange-brown to yellowish-brown, the dorsum dusted with pale grey wax mak-
ing a pattern of cross-bands; BL 1.8-3.9 mm. Most commonly on Cupressus
spp. but also on other Cupressaceae, feeding on smaller twigs in the foliated
parts of the crown and frequently causing branch die-back (Inserra et al.,
1979). It is a vector of Seiridium cardinale (Sutton and Gibson, 1972). In
Europe, southwest Asia, India (Agarwala and Raychaudhuri, 1982), North
America, and recently introduced into Africa (Mills, 1990) and Colombia,
South America (BMNH colln, leg. A. Lopez). Gunkel (1963a, b) studied
biology, ecology and natural enemies (including an Aphidius sp.) in Germany,
and Middle Eastern populations were studied in Jordan (Mustafa, 1987) and
Israel (Mendel and Golan, 1983). Oviparae and alate males in October in
Europe, but anholocycly is prevalent in regions with mild winters. Ciesla
(1992) provided an account with special reference to Africa. Some of the
literature records were based on misidentified C. fresai or C. tujafilina.
2n = 12.
628 Cinara

Cinara curtihirsuta Hottes Apterae have orange-red head and thorax and
shiny black abdomen (Voegtlin, 1976); BL 3.0-3.9 mm. On Abies concolor in
California and Oregon, USA, on branches, or on trunks of young trees,
Oviparae in September-December (Voegtlin, 1976), males unrecorded.

Cinara curvipes (Patch) Apterae are entirely shiny or dull black, in early
summer sometimes developing a covering of pale grey wax (Voegtlin, 1976);
BL 3.4-5.5 mm. On trunk or branches of Abies spp. and also feeding on
Cedrus deodora; records from Picea spp. are questionable, although Voegtlin
(1976) found a small colony on Pinus contorta. Widely distributed in USA,
Canada and Mexico (Ortiz, 1982). Oviparae and alate males in September-
October.

Cinara (Cupressobium) dahurica Szelegiewicz and Holman Apterae shining

brownish-yellow, without wax: BL 3.9-4.3 mm. Collected from twigs of
Juniperus pseudosabina in Mongolia (Szelegiewicz and Holman, 1980).
Biology and other morphs not known.

Cinara diabola Hottes Appearance in life unknown, probably dark-

coloured: BL about 3.4mm. Described from apterae and alatae collected on
Pinus coulteri in California (Hottes, 1961b) and not since recorded.

Cinara dubia Hottes and Essig Appearance in life unknown, probably dark-
coloured; BL 3.0-3.5 mm (Voegtlin, 1976). Only the aptera is described
(Hottes and Essig, 1954b), from Pseudotsuga taxifolia in Oregon, USA. Also
recorded from California (Voegtlin and Bridges, 1988). Probably a synonym
of C. pseudotaxifoliae (q.v.).

Cinara eastopi Pintera Apterae brownish; BL 4.0-5.0 mm. On Pinus

griffithii, 'under bark' (BMNH colln, leg. S.N. Chatterjee), in northwest
India. Biology and sexuales unknown. Vagrant alatae have been found on
several different plants and on snow, suggesting that this species may be com-
mon but often overlooked on pine. It is very similar to, and may prove to
be synonymous with, C. orientalis.

Cinara edulis (Wilson) Apterae gun-metal or black, often with a narrow

spinal stripe and paired patches of whitish wax powder on thorax and
abdomen, legs bicoloured black and orange (Palmer, 1926); BL 2.5-4.0 mm.
Hottes (1960b) discussed colour variation. In large colonies on bark of
branches and older twigs, or on young tree-trunks, of Pinus edulis and
P. monophylla in western USA. Oviparae and males in September-October,
eggs laid along the needles (Palmer, 1926).

Cinara engelmanniensis Gillette and Palmer 'Light cinnamon-brown to tan

or pinkish-buff, with a pair of longitudinal rows of green or black spots'
(Gillette and Palmer, 1925); BL 2.2-2.7 mm. Scattered or in small groups on
bark of branches or twigs of Picea engelmannii, usually close to the ground
 Cinara 629

(Hottes, 1961b). In western USA. Oviparae and alate males in September-

October.
Cinara essigi Hottes Appearance in life unknown; BL of aptera c. 4.5 mm.
Described from Pinus attenuata in California (Hottes, 196la). Biology and
sexuales unknown. Apart from its larger size this aphid is, to judge from
its description, closely related to and possibly synonymous with either
C. cronartii or C. newelli.
Cinara etsuhoe Inouye Apterae elongate oval, dark brown; BL 4.5-5.8 mm.
Collected on bark of trunk of Pinus densiflora in Japan, near the ground and
often in ant shelters (Inouye, 1970). Life cycle and sexuales unknown. Very
similar to and perhaps conspecific with C. sorini.

Cinara ferrisi (Swain) Apterae rather elongate oval, brown mottled with
black, slightly pruinose; BL 4.3-6.5 mm. On Pinus albicaulis and P. monticola
in western USA and British Columbia. The only recorded feeding site is the
roots (Bradley, 1961). Life cycle and sexuales unknown. Records from Abies
concolor should probably be referred to other species, e.g. C. curvipes.

Cinara flexilis Gillette and Palmer Apterae brownish-black, dull or shining,

without wax; BL 4.0-5.0 mm. Scattered on bark of twigs and small branches
of Pinus flexilis and P. strobiformis in Arizona (BMNH colln; leg. F.C.
Hottes) and Colorado, USA. Oviparae in September in Colorado (Palmer,
1952).

Cinara formosana (Takahashi) Apterae broadly oval, yellowish-brown with

dusky yellow legs mottled with dark brown; BL 4.0-5.5 mm. Immatures are
somewhat greenish (Takahashi, 1924a). On bark of one- or two-year-old
shaded branches (Inouye, 1970) of Pinus spp. (especially massoniana,
thunbergii) in East Asia (Japan, Korea, China, Hong Kong, Taiwan).
Oviparae and alate males in November (BMNH colln). C. pinitabulaeformis
Zhang, 1989 (in Zhang and Zhong, 1989) is a synonym. 2n = 10.

Cinara fornacula Hottes Apterae are pale, translucent green, or sometimes

pale brown, with slight pruinosity; BL 3.1-4.5 mm. On small branches and
current year's growth of Picea spp., living at the needle bases, not attended
by ants (see Johnson and Lyon, 1988, p. 83). Widely distributed in North
America. Oviparae and alate males collected in late August, eggs laid on the
needles (Hottes, 1933). 2n = 10.
Cinara (Cupressobium) fresai Blanchard Apterae pinkish-grey to dark
brownish-grey, dusted with white wax intersegmentally, especially on sides of
thorax, and with paired black patches on thoracic and anterior abdominal
tergites diverging in an inverted 'V' (Carter and Maslen, 1982); BL 2.2-
4.2 mm. Alatae are unusual in having the radial sector not reaching the apex
of the forewing. On foliage and adjacent woody shoots and branches of
various Cupressaceae (Juniperus spp., Cupressus spp. - see Eastop, 1972),
630 Cinara

and also recorded from Cryptomeria japonica (Taxodiaceae; Bradley, 1965b,

as C. maui). Known distribution is remarkably disjunct; England, Spain,
USA, Central and South America, Australia, New Zealand, Japan (BMNH
colln; leg. R. Takahashi), and most recently recorded from Israel (Mendel and
Zehavi, 1987). Cinara fresai is evidently anholocyclic; no sexual morphs have
been recorded, and the karyotype is structurally heterozygous (2n = 13).

Cinara gentneri Hottes Apterae light yellowish-brown to brown (Hottes,

1957a): BL 5.5-5.8 mm. On Abies grandis and A. concolor, only recorded
from Oregon, USA. Oviparae in October. Alate viviparae and males, and
biology, not known.

Cinara glabra (Gillette and Palmer) Apterae dusky olive green, olive-brown
or black, without dorsal wax; BL 2.5-3.0 mm. Singly or in small groups on
twigs at bases of needles of Pinus ponderosa in western USA; Colorado,
Arizona and Wisconsin (BMNH colln; leg. F.C. Hottes). Oviparae and alate
males in October (Gillette and Palmer, 1924, 1931).

Cinara glacialis Hottes Appearance in life not recorded; BL of aptera

c. 2.7 mm. Other morphs and life cycle unknown. Collected on Picea sitchensis
in Alaska (Hottes, 1964). Closely related to, if not synonymous with, C.
glehna.

Cinara glehna (Essig) Apterae reddish-brown to dusky brown, but in life

covered with white wax (Essig, 1915); BL 2.9-4.0 mm. On twigs of Picea spp.,
feeding on bark at bases of needles. First described heavily infesting the East
Asian species P. glehna in California, but since reported mainly from
P. pungens, which is presumably the native host. In western North America.
Sexuales and life cycle unknown. Closely related to C. braggii, and possibly
the two are variants of a single species. Cinara mariana, described from Picea
mariana in Saskatchewan, Canada, and not found since; keys to C. glehna
on the basis of the original description (Bradley, 1956b), and the only clear
difference seems to be its smaller size (BL c. 2.0mm). Cinara yukona Hottes,
described from one alate vivipara on P. glauca in Alaska (Hottes, 1964), is
also very similar and may be a synonym, as may C. glacialis.

Cinara gracilis (Wilson) Apterae are shiny black, sometimes with a whitish
wax spinal stripe on thorax and ABD TERG 1-2; BL 2.7-3.5 mm. It lives in
dense colonies on bark among needles, often near the bases of twigs and small
branches and is easily disturbed when approached (Pepper and Tissot, 1973).
The normal host is Pinus virginiana, and it is also found on P. taeda (Patti
and Fox, 1981), but the more northerly records from P. banksiana (e.g.
Bradley and Hinks, 1968) seem to be misidentifications of canatra and/or
nigra. In eastern USA.

Cinara grande Hottes Appearance in life unknown; BL of aptera 4.8-

5.1 mm. On trunk of young Abies concolor in Arizona, USA, not collected
 Cinara 631

since the original description (Hottes, 1956f), although the earlier records
of Lachnus abietis on A. concolor in California by Davidson, referred to
C. ferrisi by Swain (1918), possibly apply to this species. Other morphs and
life cycle unknown.

Cinara guadarramae Mimeur Apterae dark brown, often with two dorsal
longitudinal rows of small black sclerites; BL 2.3-3.7 mm. On young branches
of Pinus sylvestris and P. pinaster in Spain. Very similar to C. pini; a detailed
morphometric comparison is made by Garcia Sanchez and Nieto Nafria
(1978). Sexuales and life cycle are not known.

Cinara harmonia Hottes Apterae dark greenish-brown or brownish (Hottes,

1958h), with broad but fairly low black siphuncular cones; BL 2.9-4.1 mm.
Usually in colonies, feeding at needle bases on two- to three-year-old wood
of Pinus resinosa, in eastern USA and Canada. Oviparae in October, males
unrecorded (Pepper and Tissot, 1973).

Cinara hattorii Kono and Inouye Apterae dark brown to black, rather
elongate oval; BL 3.2-4.4mm. Alatae have forewing membrane pigmented
between Cula and Cu1b. On Abies sachalinensis and A. mariesii in Japan,
forming colonies on bark of trunk or old twigs l m or less above ground.
Oviparae and alate males in October-November. Cinara konoi is a synonym
(Inouye, 1956).

Cinara hirsuta Hottes and Essig Appearance in life unknown; BL of aptera

2.9-3.3 mm. Only known from original collection of apterae on Pinus mon-
ticola in Oregon, USA (Hottes and Essig, 1954a). Cinara kuchea is possibly
a synonym.

Cinara horii Inouye Apterae rather elongate oval, with yellow head, light
green thorax and green abdomen (Inouye, 1956); BL 4.4-5.3 mm. Resembling
C. pilicornis, but without a dark cross-band on ABD TERG 8. On twigs
and young shoots of Picea jezoensis in Japan and Sakhalin (Inouye, 1970).
Eggs and fundatrices have been observed in April-May, but sexuales are
unrecorded.

Cinara hottesi (Gillette and Palmer) Apterae dull bluish-black with mainly
yellow appendages (Gillette and Palmer, 1924); BL 3.0-3.5 mm. In large dense
colonies on bark of twigs of Picea spp., moving to larger branches or trunk
as season progresses (Bradley, 1961). Widespread in North America. Oviparae
and apterous males in September-October in Colorado (Hottes, 1955a).
Cinara alaskana, described from Picea sp. in Alaska (Hottes, 1964), is prob-
ably a synonym; its BL is recorded as 2.2 mm, but this is possibly a misprint.

Cinara hyperophila (Koch) (= diversiseta Börner) Aptera dark brown to

black, shining, with legs mainly or wholly dark; BL 2.7-3.6 mm. On one- to
three-year-old shoots of Pinus spp. (mugo, sylvestris), ant-attended, never in
632 Cinara

large colonies. In Scandinavia and Central Europe. Heikinheimo (1984)

redescribed the viviparous morphs (as diversiseta), and Danielsson (1987)
discussed the synonymy. Biology is little known; Koch (1855) collected
oviparae in September-October and found numerous males in April-May,
perhaps due to exceptional conditions (see Eastop, 1976). [Populations on
P. mugo that key to hyperophila may be referable to C. setosa Börner, recently
redescribed and judged to be a good species by Binazzi (1991b).] 2n = 10.

Cinara indica Verma Described from one aptera from Cedrus deodora at
Jammu, India (Verma, 1970b), possibly a small aberrant specimen of C. con-
finis. The much larger alatae described under this name by A.K. Ghosh
(1982b) are probably confinis.

Cinara inscripta Hottes and Essig Fundatrices have dark brown head and
thorax and shiny amber-brown abdomen with large prominent dark brown
siphuncular cones (Bradley, 1961), appearance of later generations in life
unrecorded; BL 3.7-4.2 mm. On Pinus albicaulis in Oregon, USA and British
Columbia, Canada, colonizing young branches in spring; otherwise biology
unknown, and sexuales are undescribed.

Cinara intermedia (Pašek) Apterae mainly pale brown, with two brownish
or greenish-black dorsal longitudinal stripes, converging on ABD TERG 5-8
(Heinze, 1962); BL 2.5-2.7 mm. On bark of trunk of young Pinus sylvestris,
as yet only collected (twice) as single apterae within colonies of C. nuda
(Pintera, 1966). Life cycle and other morphs unknown.

Cinara jucunda Hottes Appearance in life unknown; BL of aptera 2.7-

3.0mm. Apterae described from either Picea pungens or P. glauca (or both;
Hottes, 1958g) in North Dakota, USA and also recorded from Colorado and
South Dakota (Voegtlin and Bridges, 1988). Other morphs and biology
unknown.

Cinara juniperensis (Gillette and Palmer) Apterous viviparae undescribed

(keyed according to oviparae); BL 3.5-4.0 mm. On bark of tender twigs of
Juniperus communis in Colorado and Oregon, USA. Oviparae (mainly
yellowish to greenish-brown with slight pulverulence and dark tibiae) were col-
lected in late August (Gillette and Palmer, 1931). Life cycle not studied.

Cinara (Cupressobium) juniperi De Geer (= sibiricae Gillette and Palmer)

Apterae and alatae pinkish-brown with light wax dusting, not forming a
pattern, and dark legs; BL 2.2-3.4 mm. On undersides of young shoots of
Juniperus communis and its varieties, feeding on the needles; occasionally on
other Juniperus spp., but most records from other hosts are likely to be
misidentifications of tujafilina, fresai or cupressi. In Europe (including
Iceland), the Middle East (Turkey, Lebanon), Australia, New Zealand,
western USA and Canada (Ontario). Oviparae and apterous males occur in
September-October in Colorado, USA (Palmer, 1952, as sibiricae) and De
 Cinara 633

Geer (1773) recorded apterous males in Sweden. Apparently anholocyclic in

UK (Eastop, 1972) and in milder climates generally. Alatae occur mainly in
May-June. 2n = 12.

Cinara juniperivora (Wilson) Apterae are dull dusky to cinnamon or

greyish-brown with a waxy bloom, with legs mainly dark; BL 2.2-3.0 mm. On
Juniperus virginiana and J. silicicola in eastern USA, living under bark on
trunk or branches, often in ant shelters of Crematogaster - see account by
Hottes and Frison (1931; as C. difficilis). Sexuales and life cycle unknown.

Cinara keteleeriae Zhang Described on the basis of a single aptera (BL

about 3.5 mm) and two immatures collected on Keteleeria fortunei in Yunnan
Prov., China (Zhang and Zhong, 1985e). Appearance in life unrecorded.

Cinara kiusa Hottes Appearance in life not recorded. Not collected since the
original description; BL of aptera 3.2-3.7 mm, other morphs unknown. On
Abies concolor in Oregon, USA.

Cinara kochiana Börner Apterae greyish-brown to dark grey, slightly wax-

powdered, with a darker pattern of small sclerites and often a spinal stripe;
BL 4.7-6.1 mm. On Larix spp., forming ant-attended colonies in bark crevices
on lower part of trunk or bases of older branches, or (especially in mid-
summer) on large partly exposed roots. Recorded from most parts of Europe
except the Iberian peninsula, but rather rare (Eastop, 1972). Oviparae and
males in October-November, eggs laid in bark crevices. Kloft et al. (1960)
summarize biological information and Pontin (1960) found a heavily para-
sitized population on larch roots. 2n = 10. In Japan and Korea, similar aphids
have longer antennal hairs and are regarded as a subspecies, C. kochiana kochi
(Inouye, 1962, 1970).

Cinara kuchea Hottes Appearance in life not recorded; BL of paratype

apterae 2.3-2.6 mm. Described from Pinus monticola in Idaho, USA (Hottes,
1958e) and similar but larger aphids (BL up to 3.5mm) have been collected
in British Columbia, Canada; 'Cinara sp. 3' of Bradley (1961) is possibly this
species. Except for differences in hair length, C. kuchea resembles and may
prove to be synonymous with another little-known species on P. monticola,
C. hirsuta (q.v.). Biology unknown. (Bradley's species formed large, dense
colonies on small branches and roots of young trees.)

Cinara lachnirostris Hille Ris Lambers Apterae darkish-brown; BL 2.4-

2.8mm. On Pinus griffithii in Pakistan, Nepal and Kashmir (BMNH colln).
Feeding site not recorded and life cycle unknown (A.K. Ghosh, 1982b).
Bhagat (1981) reared a parasitoid (Pauesia pini) from this aphid. 2n = 8
(J. Dutta and D.C. Gautam, pers. comm.).

Cinara (Cedrobium) laportei (Remaudière) Apterae are broadly oval and

dorsoventrally flattened, pale brown and grey with a narrow pale spinal stripe
634 Cinara

from head to anterior abdomen; BL 1.5-2.0 mm. In small dense colonies on

twigs and on small shoots of lower branches of Cedrus spp. in Europe, North
Africa (Algeria, Morocco) and South Africa. Its range currently seems to be
expanding. Immatures probably of this species were also collected in Himachal
Pradesh, India (BMNH colln, leg. A. Chowdhury). Sexuales are produced in
October at high altitudes in Morocco, but in many places this aphid is prob-
ably anholocylic. Biological control with a specific parasitoid from Morocco
was undertaken in southern France (Fabre and Rabasse, 1987). Remaudière
(1954) erected the genus Cedrobium for this species which differs from
other Cinara in its 5-segmented antennae, strongly sclerotic tergum and
curious, mace-like dorsal hairs. However on phylogenetic grounds and taking
into account recent biochemical evidence (Lampel and Burgener, 1987),
Cedrobium should clearly be treated as a subgenus within Cinara.

Cinara laricifex (Fitch) Apterae dark brown to bronze, without wax; BL

2.5-3.5 mm. On Larix laricina, in small numbers on new growth near tips of
twigs, or later also forming colonies on branches or trunk (Bradley, 1959).
Males are alate, and eggs are laid on the bark (Bradley, 1961). In Canada and
northeastern USA. This species has sometimes been confused in the North
American literature with the European C. laricis (Hartig) - see Hottes (1953a).
2n = 10.

Cinara laricifoliae (Wilson) Apterae dark greenish-brown to black with a

dusting of white wax and a pale spinal stripe (Wilson, 1915; Bradley, 1961);
BL 4.2-4.6 mm. On Larix occidentalis, in small colonies on small and
medium-sized branches, feeding at bases of needle clusters. Recorded from
Oregon, USA and British Columbia, Canada. Life cycle unknown.

Cinara laricionis Binazzi Apterae dark bronze; BL 2.8-3.7 mm. In colonies

on young growth and one-year-old branches of Pinus nigra var. maritima
(= P. laricio) in Italy and Corsica (Binazzi, 1980a, b), and apparently specific
to this host. Very similar to C. maritimae. Oviparae and alate males in
October-December (Binazzi, 1983c).

Cinara laricis (Hartig) Apterae dark greyish-brown speckled with black,

usually with a wax bloom; BL 3.0-5.1 mm. In small dense colonies on twigs
of lower branches, or trunks of young larch trees (decidua, leptolepis,
x eurolepis, gmelinii). Throughout Europe and also recorded from Mongolia
(Szelegiewicz, 1969), China (Fang et al., 1983) and Japan (Inouye, 1962). The
copious honeydew crystallizes to form the 'Lärchenmanne' which is highly
regarded by beekeepers in Central Europe (e.g. Manino et al., 1985).

Cinara longipennis Matsumura Apterae dark brown to black with dark legs,
body dusted with wax laterally and ventrally (Moritsu, 1983); BL 6.0-7.0 mm.
On Abies spp. (firma, holophylla, sachalinensis), in large spring colonies with
abundant alatae on trunk and older branches of young trees, apterae moving
down onto the roots in summer (Inouye, 1970). In Japan and Korea. Oviparae
 Cinara 635

in Korea in late October (Paik, 1972; but the aphid illustrated by Paik as
longipennis is much paler, yellow-brown with black markings).

Cinara longirostris Börner Appearance in life unknown; BL of aptera

c. 4.7 mm. Only known from type specimens from Pinus sylvestris in French
Alps, which have recently been redescribed by Binazzi (1990) and judged to
be a good species. The available material (one aptera and one ovipara) is
inadequate for inclusion in a key. Resembling C. pini, except for large size
and absence of a mesosternal tubercle.

Cinara (Cupressobium) louisianensis Boudreaux Apterae green, with a

dorsal pattern of white wax (Boudreaux, 1949); BL 1.7-2.2 mm. On branches
of Thuja sp. in southern USA and on a Cupressus sp. in Mexico (Voegtlin
et al., 1986). Recorded also from Taiwan (Tao, 1958, on Thuja orientalis);
Korea (Paik, 1972); Tasmania (Eastop, 1966) and New Zealand (BMNH colln,
leg. VFE; on Th. occidentalis and Chamaecyparis lawsoniana). Biology and
sexuales unknown. 2n = 12.

Cinara lyallii Bradley Apterae dark brown to dull black, dusted with grey
wax powder; BL c. 4.0 mm. In small loose colonies on upper sides of twigs
of previous year's growth of Larix lyalli at high elevations (above 2000 feet)
in Alberta, Canada. Oviparae and both apterous and alate males are recorded,
and there is possibly an abbreviated life cycle (Bradley, 1961). Closely related
to C. laricifoliae.

Cinara maculipes Hille Ris Lambers Apterae pale brown with legs, espe-
cially femora, mottled with dark brown in pantherine fashion; BL 3.7-
4.9mm. On Pinus griffithii in northern India (Himachal Pradesh, Uttar
Pradesh, Jammu, Kashmir) and Pakistan (Muree). Feeding site not recorded.
Oviparae and apterous males in December (Chakrabarti and Raha, 1988). Das
and Chakrabarti (1989a) reared a new Pauesia sp. from this aphid. 2n = 12
(Kurl and Chauhan, 1986).

Cinara maghrebica Mimeur Apterae chocolate brown with a dorsal pattern

of white wax dust (Mimeur, 1934); BL 2.0-2.9 mm. Living, often in dense
colonies, on young twigs of Pinus spp. (canariensis, halepensis, pinaster,
pinea). In the Mediterranean area (Italy, France, Spain, Morocco) and also
recorded from Argentina (Delfino and Eastop, 1982). Garcia Sanchez and
Nieto Nafria (1978) provide a detailed morphological comparison with the
closely-related C. maritimae. Binazzi (1983a) differentiates populations on
P. halepensis in Italy as a subspecies, C. maghrebica garganica. Cinara
pasheki Pintera is a synonym. 2n = 10*.

Cinara manitobensis Bradley Apterae reddish-brown covered with dense

grey wax powder; BL 2.4-3.2mm. Described from Juniperus horizontalis
in Manitoba, Canada. Oviparae and apterous males in October (Bradley,
1963).
636 Cinara

Cinara maritimae (Dufour) (= excelsae Hille Ris Lambers), Plate 16a

Apterae brownish, covered in greyish wax dust; BL 2.6-4.1mm. Living
separately or in small groups among needle bases on young twigs of Pinus
spp., with a relatively wide range of hosts in subsect. Pinus. In the Mediterra-
nean area and Middle East, and introduced into South America (Argentina,
Brazil, Chile). Oviparae and alate males in October in Italy (Binazzi, 1978).
Anholocyclic overwintering probably occurs frequently. Mustafa (1987)
studied population biology in Jordan and Michelena Saval and Gonzalez
Funes (1988a) recorded parasitoids (Pauesia spp.) in Spain. Probably confused
in earlier literature with C. pinea (Eastop, 1976). 2n = 16*.

Cinara matsumurana Hille Ris Lambers (= abietis Matsumura) Apterae

with greenish-brown head and thorax and green abdomen; BL 2.3-3.0 mm.
On young twigs and branches of Abies spp. (homolepis, sachalinensis) in
Japan, feeding among needles, usually on upper parts of young trees.
Oviparae and alate males in October (Inouye, 1970). 2n = 10.

Cinara medispinosa (Gillette and Palmer) Apterae light cinnamon-brown

to dark metallic brown, with a pattern of black dorsal sclerotic markings
(may not be evident in midsummer populations), and some wax powder
spinally and laterally (Palmer, 1952); BL 3.0-4.0 mm. On bark of tender twigs
and young branches of Pinus contorta and its varieties (plus one record from
P. albicaulis), in western USA and Canada. Oviparae and males in October.
Closely related to brevispinosa and murrayanae; the taxonomy of this group
of species requires clarification.

Cinara melaina Boudreaux Apterae are shiny dark brown, almost black; BL
2.8-3.3 mm. Normally apterae have an uninterrupted cross-band across ABD
TERG 8, whereas in C. atlantica, with which it may form mixed colonies, this
band is usually interrupted medially. On current and previous year's growth
of Pinus spp. of subsect. Australes, feeding at needle bases; there are also
single records from P. resinosa and P. virginiana (Pepper and Tissot, 1973).
In southeastern USA, north to Delaware. Males unrecorded, and oviparae are
rare; probably anholocyclic in southern states. Patti and Fox (1981) studied
populations on P. taeda in South Carolina. Very similar when slide-mounted
to the western US species, C. ponderosae.

Cinara metalica Hottes Apterae are mainly cinnamon-brown with a pinkish

cast, covered by a mottling of wax powder that is thickest on dorsal abdomen
anterior to siphunculi, but with siphunculi and the dorsal cuticle posterior
to them conspicuously bronze-coloured and shining (Hottes, 1960b); BL
3.3-3.6 mm. On Pinus edulis, feeding among needles on the younger branches
of young healthy trees with greenish bark. In Arizona and Colorado, USA.
Sexuales in October; the three known male specimens all have vestigial wings.

Cinara minoripinihabitans Zhang Body of aptera with a thick covering of

wax powder, head and thorax pale, legs black (Zhang and Zhong, 1989); BL
 Cinara 637

c. 2.4mm. On Pinus koraiensis in Heilongjiang Prov., China. Other morphs

and life cycle unknown.

Cinara minuta Hottes and Knowlton Appearance in life unknown, perhaps

rather pale; BL (ovipara) 2.3-2.4 mm. Only known from the oviparous
morph, collected on Abies lasiocarpa in Utah, USA (Hottes and Knowlton,
1954).

Cinara moketa Hottes Apterae shining dark brown with cross-bands of grey
pulverulence; BL 3.3-4.1 mm (Voegtlin, 1976). In colonies on branches, or on
trunks of young trees, of Pinus lambertiana in California and Washington,
USA. Sexuales unknown, apparently anholocyclic in California (Voegtlin,
1976). Tilles (1984) studied ant attendance, and predation by an elaterid
beetle. Cinara hirticula Hottes is probably a synonym.

Cinara mongolica Szelegiewicz and Holman Apterae brownish, without

wax; BL 3.7-3.9 mm. Type specimens were from a small colony, not attended
by ants, on young twigs of Pinus sibirica in Mongolia (Szelegiewicz and
Holman, 1980). Oviparae and alate males were collected in August. Very
similar to C. cembrae, and possibly all records of C. cembrae in the Far East
(e.g. Inouye, 1970; Pashchenko, 1988b) should be referred to mongolica, in
which case P. cembra, P. pumila and P. koraiensis are additional hosts.

Cinara montanensis (Wilson) Appearance in life unknown, legs almost

entirely black; BL c. 4.0mm. Known only from the original collection of
alatae and apterae on Pinus 'laricis'( = P. nigra var.?) in Massachusetts, USA
(Wilson, 1919b).

Cinara montanesa Hottes Appearance in life unknown; BL of aptera

c. 5.7mm (but that of alata is given as only c. 3.5mm). Known only from
original collection of apterae and alatae on Pinus coulteri in California, USA
(Hottes, 1961c).

Cinara (Cupressobium) mordvilkoi Pašek Apterae have light coffee-brown

wax-dusted head and thorax and darker shining bronze abdomen; BL 2.7-
3.3mm. On young growth and branches of Juniperus communis in
Czechoslovakia (Pasek, 1954), and also recorded from Poland (Szelegiewicz,
1962) and Latvia (Rupais, 1961). Specimens from roots of Juniperus on the
island of Öland, Sweden, attended by Lasius flavus, appear to be this species
(BMNH colln, leg. VFE). Holocyclic; males are apterous (cf. C. cupressi).
Klimaszewski et al. (1977) described biochemical differences from cupressi.

Cinara murrayanae (Gillette and Palmer) Apterae yellow-brown to dark

brown; BL 3.0-4.0 mm. On bark of twigs and small branches of Pinus con-
torta and its varieties, in western North America; records from other hosts
are probably misidentifications of other species. Oviparae and alate males in
October (Gillette and Palmer, 1924). Difficult to distinguish from less sclerotic
638 Cinara

midsummer individuals of C. medispinosa and resembling C. atlantica except

in length and shape of hairs.

Cinara nepticula Hottes Only known from alate viviparae, BL 3.4-3.6mm,

collected on Picea rubens in Nova Scotia, Canada (Hottes, 1958c, 1961b).
Probably closely related to, if not synonymous with, C. braggii and/or
C. glehna.

Cinara neubergi (Arnhart) Apterae of variable colour, light to dark brown,

with a slight dusting of wax (Arnhart, 1930); BL 4.0-5.0 mm. On bark of twigs
of Pinus mugo, feeding among needle bases, in mountainous regions of
Central Europe (Austria, Bulgaria, Germany, Italy, Switzerland, Ukraine).
Oviparae in September (BMNH colln); males unknown (Pintera, 1966). A
member of the C. pinea group.

Cinara newelli Tissot Apterae yellowish-brown to dark brown with a cover-

ing of grey wax (Tissot, 1939); BL 2.7-3.1 mm. Like C. cronartii, this species
feeds at rust lesions (Cronartium fusiforme) on Pinus spp. (elliotti, taeda).
Only known from Florida, USA; sexuales and life cycle not recorded (Pepper
and Tissot, 1973).

Cinara nigra (Wilson) Apterae shining chocolate brown, without wax; BL

2.5-3.0 mm. In large dense colonies on undersides of branches, or on trunk
of young trees, of Pinus banksiana and P. contorta. In northwestern USA east
to Michigan and in Canada from British Columbia east to Ontario. Apterous
males and oviparae in late September (Pepper and Tissot, 1973). Eggs are laid
in the cleft between two needles just above the sheath (Bradley, 1961, as
canatra). Foottit and Mackauer (1990) studied morphological variation within
and between populations on P. contorta.

Cinara nigripes Bradley Apterae dark brown to bluish-black, with scattered

white wax (Bradley, 1962); BL 2.8-3.4 mm. In dense colonies on trunk of
young trees of Picea spp. (glauca, mariana, sitchensis) in Canada (British
Columbia, Ontario) and Alaska. Oviparae and small apterous males in mid-
August in Ontario.

Cinara nigrita Hottes and Essig Only known from a single alata, distin-
guished by its uniformly dark wings, collected in Arizona, USA. The host is
unknown but surmised to be Pinus ponderosa (Hottes and Essig, 1953b).

Cinara nimbata Hottes Apterae have black head and thorax dusted with
grey wax and abdomen grey-green dorsally with lateral areas cinnamon-brown
(Hottes, 1954c); BL 3.7-4.2 mm. The legs are characteristically patterned, with
a contrasting pale band across the dark distal part of each femur and tibiae
spotted with black. Alatae have diffuse patches of fuscous on forewings as
in C. costata, to which it is closely related. Living singly particularly on stubby
branches close to trunk of tree and attended by ants (Hottes, 1954c). On Picea
 Cinara 639

engelmanni in western North America from Colorado to Alaska; an ovipara

from an unknown host in Labrador and an alata collected on P. mariana in
Quebec (BMNH colln, leg. F.C. Hottes) also seem to be this species. Oviparae
and alate males in September in Colorado (Hottes, 1954c).

Cinara nitidula Hottes Apterae various shades of brown to almost black,

shining, wax-dusted only on head and sides of thorax; BL 2.0-2.3 mm. On
Pinus edulis, apparently restricted to older terminal branches of mature trees,
on yellow bark among scales (Hottes, 1960b). In Colorado and Utah, USA.
Apterous males and oviparae in October (Hottes, 1956e).

Cinara nuda (Mordvilko) (= escherichi Börner) Apterae are brownish to

bronze, shining with little or no wax; BL 3.5-4.3 mm. In large colonies on
Pinus sylvestris, on trunk and basal parts of older branches of young trees,
or on two- to eight-year-old parts of leading shoots of older trees. In northern
and Central Europe (England, Finland, Poland, Germany, Czechoslovakia,
Austria). Apterous males and oviparae occur in Central Europe in August-
October. Scheurer (197la) studied populations over a three-year period in
Germany. Danielsson (1987) established the correct name for this species.
2n = 10.

Cinara obscura Bradley Apterae dark brown to black, with pale sections on
tibiae; BL c. 2.5 mm. In large dense colonies on bark of small branches of
Picea glauca in Canada, from Alberta to Newfoundland (Bradley 1953, 1961)
and also recorded from P. engelmannii in British Columbia. Fundatrices were
found on P. glauca in Ontario (BMNH colln, leg. Bradley), but sexuales are
not recorded. C, pallidipes Hottes, described from P. glauca in Maine, USA,
may be a synonym.

Cinara occidentalis (Davidson) Apterae golden-brown with bluish-white

wax either in cross-bands or forming a flocculent covering of body and appen-
dages; BL 2.1-3.2 mm. On bark of one-year-old twigs of Abies spp. in western
USA and Canada (British Columbia, Quebec). Oviparae and alate males in
September-November. Ant attendance was studied by Tilles and Wood
(1986). Heavy parasitization was noted by Knowlton (1930). (N.B. An Indian
Abies-feeder, C. tistaensis (q.v.), would probably key out to occidentalis,
although clearly not this species.)

Cinara ontarioensis Bradley Apterae dark brown, shining, without wax; BL

2.4-2.8 mm. On Pinus banksiana in Canada (Ontario, Manitoba), feeding on
needle bases of one- and two-year-old branches and causing yellowing of
needles in late spring and summer. Numerous alatae are produced in the
second generation. Oviparae and alate males in autumn (Bradley, 1962).

Cinara oregonensis (Wilson) Apterae light brick-red to rust-red, shining; BL

2.5-3.0 mm. On Pinus contorta and P. ponderosa and their varieties in
western USA and Canada, living among scales of young cones. The second
640 Cinara

generation are nearly all alate and fly to new cones. Oviparae and apterous
males in September-October; eggs are laid on undersides of scales of new
cones.

Cinara oregoni Hottes and Essig Appearance in life not recorded; BL of

aptera 4.0-4.4 mm (Hottes and Essig, 1953a). On Pinus albicaulis in Oregon,
USA and also recorded from this host in British Columbia, Canada. Other
morphs and biology unknown.

Cinara orientalis (Takahashi) Apterae blackish-brown, shining, not waxy;

BL 4.5-5.0 mm. Described from Pinus sp. in Taiwan (Takahashi, 1925); in
Japan it feeds on branches of Pinus densiflora, in ant shelters constructed by
Lasius niger (Inouye, 1970). Also recorded from Korea (Paik, 1972) and there
are specimens in the BMNH collection from China (on P. armandii) and Nepal
(one alata). Sexuales and life cycle unknown. Zhang in Zhang and Zhong
(1985e) described a subspecies, P. orientalis lijiangensis, from P. yunnanensis
in China, based on differences in antennal sensoriation that are probably
within the normal range of the species. Cinara eastopi in India may also prove
to be a synonym of orientalis.

Cinara osborni Knowlton Apterae buff-coloured to grey, dusted with wax,

with four black spots on thorax (Hottes, 1960c); BL 3.3-4.2 mm. Living
solitarily on the twigs of Abies spp. (concolor, lasiocarpa) in western USA.
(Originally described from Pseudotsuga (Knowlton, 1942), but this is unlikely
to be a normal host.) Sexuales and life cycle unknown.

Cinara pacifica (Wilson) Only known from original collection of three

alatae and 11 immatures from Abies grandis in northern California. Distin-
guished by the unusual siphunculi which are flattened and inconspicuous,
without hairs or pigmented conical base (Palmer, 1945). Hind tibiae are
entirely dark.

Cinara palaestinensis Hille Ris Lambers Apterae chestnut-brown to

yellowish-green (Binazzi, 1978); BL 2.0-3.3. mm. In large dense ant-attended
colonies on new growth of Pinus halepensis (incl. var. brutia) in the Mediterra-
nean area and southwest Asia (Israel, Italy, Spain, Turkey). Sexuales not
recorded; perhaps mainly anholocyclic (Barbagallo and Stroyan, 1982).
Binazzi (1983a) differentiated some Italian populations as a subspecies, C.
palaestinensis apulica. 2n = 10*.

Cinara parvicornis Hottes Only the ovipara and alate male of this species
are described, collected on Pinus contorta in Montana in September. BL of
ovipara 2.5-3.2 mm (Hottes, 1958d).

Cinara paxilla Zhang Appearance in life unrecorded; BL of aptera

c. 4.4mm. On Pinus sp. in Tibet (Zhang and Zhong, 1981b). A short-haired
species, possibly close to C. formosana.
 Cinara 641

Cinara pectinatae (Nördlinger) Apterae are bright olive green with three
diffuse paler green longitudinal bands; BL 2.8-5.0 mm. On Abies spp.,
especially A. alba ( = pectinata). It occurs throughout Europe eastward to
Turkey, living singly on small branches where it sits on the needles, feeding
at the junction of petiole and stem. Oviparae and alate males in October. A
copious honeydew-producer, important to forest beekeepers in Central
Europe and its population dynamics has been much studied because of this
(e.g. Bloc, 1987, in France; Liebig, 1988, in Germany). 2n = 6* (Germany,
2 samples; but Rukavishnikov (1979) recorded 2n = 12 from the former USSR).

Cinara pergandei (Wilson) Apterae are rather globose, shiny, brown to

bronze without wax; BL 3.0-4.4 mm. Active insects living singly on twigs and
new shoots of many Pinus spp. in subsections Australes and Contortae. In
eastern USA, across Canada, and in Cuba (Hernandez and Rodriguez, 1985).
Oviparae and alate males in September-November (Pepper and Tissot, 1973).
Relations with parasitoids and ants were recorded in Manitoba (Bradley and
Hinks, 1968). 2n = 14*.

Cinara (Cupressobium) petersoni Bradley Apterae have dark greyish-brown

head and thorax and greenish-brown abdomen, and are dusted with greyish-
white wax (Bradley, 1963); BL 2.9-3.4 mm. Described from Juniperus hori-
zontalis in Manitoba, Canada; specimens conforming to this description have
also been collected in Majorca and Israel (on J. oxycedrus; BMNH colln).
Alatae are undescribed, and sexual morphs and life cycle are unknown.

Cinara piceae (Panzer) (= grossa Kaltenbach) Apterae are uniformly jet

black 'resembling the texture and shape of old droplets of tar' (Carter and
Maslen, 1982); BL 4.5-6.7 mm. Forming large colonies in spring on undersides
of older branches and on trunks of Picea spp., often moving to ground level
or roots in summer (Pintera, 1966; as grossa). Throughout Europe, and what
is nominally the same species also occurs in Siberia (Pashchenko, 1988b),
China and Japan. Numerous alatae are produced in May-June. Oviparae
appear in September-October, move to current year's growth and lay wax-
dusted eggs on needles. Males are apterous, small and usually overlooked
according to Carter and Maslen (1982), but Inouye (1970; as grossa) reported
alate males in Japan. Starý (1976) studied the natural enemy complex in
Central Europe. 2n = 10* (UK) or 12 (Rukavishnikov, 1979, European
Russia).

Cinara piceicola Cholodkovsky (= viridescens, = stroyani) Apterae have

dark brown head and thorax and pale olive-buff abdomen with two longi-
tudinal faint greyish-green stripes (Carter and Maslen, 1982); BL 2.1-4.2 mm.
On Picea spp., in colonies on bark of woody shoots between needle bases in
spring, moving to older branches and roots in summer (Börner and Franz,
1956). In northern, Western and Central Europe. Numerous alatae are pro-
duced in May-June. Oviparae and apterous males occur from July onwards
(Pintera, 1966). Kunkel (1990) studied its relationship with ants. Danielsson
642 Cinara

(1987) discussed the confused nomenclature of this species. Cinara alba

Zhang, 1982 (in Zhang and Zhong 1982b), described from P. asperata in
China, is very similar and possibly a synonym. 2n = 8*.

Cinara pilicornis (Hartig) Apterae are either orange-brown or greyish-green,

secreting dense white wax wool; BL 2.1-4.7 mm. On Picea spp., initially in
spring in small colonies on undersides of the previous year's twigs of Picea
spp., moving onto new growth after bud-burst. It may also colonize Tsuga
heterophylla (Forbes and Chan, 1976; Carter and Maslen, 1982). Throughout
Europe (incl. Iceland), east to Kazakhstan and introduced to Australia, New
Zealand, North and South America (Eastop, 1972; Carrillo, 1977; Sunde,
1984). Numerous alatae are produced in May-July, oviparae and males in
early August-November. Kettner (1985) studied populations in relation to
honeydew production in Germany, Parry (1979) studied egg survival,
Michelena Saval and Gonzalez Funes (1988a) recorded parasitoids and
Edwards (1981) described a fungal infection. 2n = 10* (in UK and New
Zealand) or 14 (Rukavishnikov, 1972, European Russia).

Cinara pilosa (Zetterstedt) (= maculata Gavrilova) Apterae are rather pale

yellowish- to reddish-brown or olive green, with spotted legs; BL 3.1-5.2 mm.
On Pinus sylvestris, feeding singly or in small groups on young shoots of older
branches of mature trees (e.g. among male cones), or on stunted twigs of
younger trees (Holopainen and Heikinheimo, 1983). In northern and Central
Europe and also recorded from Italy (Binazzi, 1988). Biology and sexual
morphs are not reliably recorded because of confusion with the closely-related
C. pinea. Danielsson (1987) discussed the nomenclature and differences from
C. pinea. 2n = 8*.

Cinara pinata Hottes Apterae have dorsal abdomen deep bronze-brown

with a yellowish spinal stripe and black markings, becoming darker with some
wax powder later in season (Hottes, 1955c); BL 2.9-3.4 mm. On older trees
of Pinus edulis, either in small colonies on bark of four- or five-year-old
branches (Hottes, 1955c), or in larger colonies on smaller terminal branches
among the needles (Hottes, 1960b). Only known from Colorado, USA.
Apterous males and oviparae in September-October.

Cinara pinea (Mordvilko) Apterae light orange-brown (spring) to grey or

dark brown (summer); BL 3.1-5.2 mm. On new shoots of Pinus sylvestris and
sometimes other Pinus spp. including nigra in dry areas. Throughout Europe,
Georgia, eastern Siberia (Pashchenko, 1988b) and introduced to North
America (see Voegtlin and Bridges, 1988). Oviparae and males in October.
Biological studies include Kidd (1985, 1990; population dynamics), Kidd and
Tozer (1984, 1985a, b; induction of alatae, overwintering survival, distribution
within canopy), Sudd (1983; ant attendance) and Michelena Saval and
Gonzalez Funes (1988a; parasitoids). 2n = 10, 11 or 14 (Blackman, 1990),
suggesting that there may be sibling species still to be recognized.
 Cinara 643

Cinara pini (Linnaeus) Apterae are dark grey or greyish-green with black
markings, and with either a slight bronze iridescence or a dusting of grey wax
(Carter and Maslen, 1982); BL 2.5-4.4 mm. On young shoots of Pinus
sylvestris in spring (Pintera, 1966), later on undersides of older foliated twigs
or branches. There are also records from many other Pinus spp., but many
of these are likely to be misidentifications or misapplications of the name pini
(Eastop, 1972). Most European records of C. pini from P. mugo should prob-
ably be referred to P. montanicola Börner. Binazzi and de Silva (1993)
distinguished this as a valid species and noted that it lived in dense colonies
preferably on two- to three-year-old often bare branches of mountain pine.
Throughout Europe and apparently also in Siberia (Pashchenko, 1988b) and
Japan (Moritsu, 1983). North American records cited as pini are probably of
pinea or a native species. Oviparae and apterous and/or alate males in
September-October. Larsson (1985) studied within-crown distribution of C.
pini, Sudd (1983) studied ant attendance, and Michelena Saval and Gonzalez
Funes (1988a) recorded parasitoids. To judge from type material recently
remounted, C. oblonga (Del Guercio) is a new synonym. 2n = 10.

Cinara pinidensiflorae (Essig and Kuwana) Apterae dark reddish-brown

with silvery wax dorsal markings (Essig and Kuwana, 1918); BL 2.3-3.5 mm.
In dense colonies on twigs and branches of oriental Pinus spp. in Japan,
Taiwan, Korea and China (BMNH colln). Oviparae and alate males in Japan
in October-November (Inouye, 1970). Cinara atratipinivora Zhang (Zhang
and Zhong, 1982b) seems to be a synonym. 2n = 10.

Cinara piniformosana (Takahashi) Apterae blackish-brown dusted with

grey wax dorsally, orange ventrally, with black legs; BL 2.8-4.2 mm. On
young trees of Pinus densiflora and P. thunbergii; in spring at bases of new
shoots, later moving to one- or two-year-old parts of branches (Inouye, 1970;
Furuie et al., 1990). In Japan, Korea, Taiwan, Siberia (Pashchenko, 1988b),
Malaya (BMNH colln), and on introduced oriental pines in Brazil (Eastop,
1976). Holocyclic in northern Japan, with alate males (Inouye, 1970), but
populations studied by Furuie et al. (1990) in western Japan were partially
anholocyclic. 2n = 10.

Cinara pinihabitans (Mordvilko) Apterae and alatae dull reddish-brown,

often dusted with grey wax; BL 3.1-4.6 mm. On bark of thin two- to eight-
year-old branches of Pinus sylvestris and also recorded from P. mugo var.
mughus. Colonies are rarely seen, possibly because they tend to occur high
in crown of tree (Stroyan, 1973). Widely distributed in Europe (Eastop, 1972).
Oviparae and alate males in October.

Cinara piniphila (Ratzeburg) Apterae greyish-brown with a rusty tinge,

dusted with wax (Heinze, 1962); BL 3.9-4.5 mm. On bark of one- or two-year-
old twigs of Pinus sylvestris, feeding among needles. In northern, Central and
southern Europe. Sexuales and life cycle not recorded. Tsinovsky and Yegina
(1977) used Entomophthora virulenta in biocontrol of this species in Latvia.
644 Cinara

Cinara piniradicis Bradley Apterae and alatae are dark brown, without wax;
BL 4.0-4.8 mm. In colonies on roots of young Pinus banksiana in central
Canada and USA (Michigan). Apterous males and oviparae in September-
October; eggs are laid singly on inner side of needles just above the follicle
sheath (Bradley and Wighton, 1959; Pepper and Tissot, 1973).

Cinara pinivora (Wilson) Apterae have shiny dark brown head, lighter
brown thorax and abdomen with dark dorsal sclerites and spots of grey wax,
black steep-sided siphuncular cones and all legs with extensive pale yellow
sections (Tissot, 1944; as osborni); BL 3.3-4.2 mm. In dense colonies at tips
of branches, or scattered along older sections of twigs, of Pinus spp. of sub-
sections Australes and Contortae in eastern USA and Canada. Oviparae and
alate males in October-November (Pepper and Tissot, 1973).

Cinara pinona Hottes Apterae have brown wax-dusted head and thorax,
dark brown or greenish-brown abdomen with a metallic sheen; BL 3.1-
3.3 mm. In large colonies on twigs of Pinus edulis among the needles, usually
on older trees with yellowish to brownish bark (Hottes, 1960b). In western
USA. Oviparae and alate males in October (Hottes, 1956f).

Cinara poketa Hottes Apterae shiny brownish-black, without wax (Hottes,

1956f); BL c. 2.6mm. On small, needle-free branches of Pinus edulis in
western USA (Arizona, Colorado). Oviparae in October (Hottes, 1960b),
males unknown.

Cinara ponderosae (Williams) Apterae have dull brown head and thorax
and shiny golden-brown abdomen, with a dorsal pattern of wax patches; BL
2.6-4.2 mm (Voegtlin, 1976). On bark usually near tips of twigs and small
branches of Pinus ponderosa and its varieties; sometimes on other Pinus spp.
in subsection Ponderosae and also recorded from planted P. canariensis and
P. roxburghii. Records from other pines may be misidentifications. In western
North America. Oviparae and alate males in October-November in Colorado
(Palmer, 1926), but populations studied in the Californian Sierra Nevada were
anholocyclic (Voegtlin and Dahsten, 1982).

Cinara pruinosa (Hartig) (= bogdanovi Mordvilko) Apterae are dark green

or brown, sometimes with a bronze metallic tinge, with blackish markings and
some lateral wax, and very prominent black siphuncular cones (Carter and
Maslen, 1982); BL 2.4-5.0 mm. In small colonies on woody twigs of Picea spp.
in spring, but later found at base of trunk and on roots in ant shelters (Pintera,
1966). Oviparae and alate males occur in September-October, but anholo-
cyclic overwintering on roots also occurs. Throughout most of Europe east-
ward to Turkey, and in North America (most records as palmerae; see
Voegtlin and Bridges, 1988). Cinara rara Bradley, described from sexuales on
Picea mariana in Saskatchewan, is also a synonym. Similar populations in
Japan (Inouye, 1970) and Siberia (Pashchenko, 1988b) tend to be shorter-
haired and are regarded as a subspecies, C. pruinosa ezoana Inouye, the
 Cinara 645

population dynamics of which was studied by Furuta and Takai (1983).

2n = 10*.

Cinara pseudoschwartzii Palmer Apterae brownish-black to dull black with

white wax markings; BL 3.3-4.0 mm. On bark of twigs of Pinus ponderosa
in Colorado, USA (Palmer, 1936a). Holocyclic (fundatrices in May), but
sexuales undescribed.

Cinara pseudotaxifoliae Palmer Apterae dark brown to black with burnt-

orange mid-dorsal area and wax powder on head, prothorax and in large
patches anterior and posterior to siphunculi (Palmer, 1952; Voegtlin, 1976);
BL 2.2-3.4 mm. On two-year-old or older twigs and branches of Pseudotsuga
menziesii in western North America. Oviparae and alate males in October-
November. Schowalter et al. (1988) studied ant attendance. Cinara dubia
Hottes and Essig, described from Oregon, is probably a synonym, and
C. vagabunda (q.v.) is possibly only a long-haired variant population.

Cinara pseudotsugae Wilson Apterae rather shiny amber to golden-brown,

sometimes with paired black dorsal markings and/or slight wax dusting
laterally and intersegmentally; BL 2.4-3.7 mm. On terminal shoots and
branches of Pseudotsuga menziesii in western North America, attended by
ants (Johnson and Lyon, 1988, p. 85). Oviparae and alate males in October
(Wilson, 1912). Cinara taxifoliae Swain is a synonym (Voegtlin, 1976). Palmer
(1926, 1952) and Hottes (1961d) apparently had a different species with darker
legs, possibly a form of pseudotaxifoliae. See also C. splendens.

Cinara puerca Hottes Apterae shiny brownish-black dorsally, sometimes

with a paler abdominal spinal stripe, and pale reddish-brown ventrally, with-
out wax (Hottes, 1954c); BL 3.0-4.4 mm. On trunk and branches of Pinus
edulis, in crevices in bark and/or in ant shelters. Recorded from Arizona and
Colorado, USA. No sexuales known and the populations studied in Colorado
were anholocyclic (Hottes, 1955a).

Cinara pulverulens Gillette and Palmer Apterae yellowish-brown with black

pleural markings, covered with dense white wax powder; BL 2.2-2.9 mm. On
bark of small twigs of Juniperus spp. in western USA (Colorado, Idaho,
Utah). Oviparae and alate males in September-October (Palmer, 1952). Seems
likely to be a synonym of C. burrilli, as it differs only in hair length, colour
and amount of wax.

Cinara radicivora Voegtlin Apterae shining dark greenish-black with scat-

tered brown patches; BL 4.4-6.1mm. On root crown of Abies concolor,
attended by Camponotus spp., in California and Oregon, USA (Voegtlin,
1982). Life cycle and sexuales unknown.

Cinara rigidae Hottes Apterae 'probably black' (Hottes, 1958a); BL

c. 2.4 mm. Known only from one aptera and two alatae collected on Pinus
646 Cinara

rigida in New Hampshire, USA; type specimens redescribed by Pepper and

Tissot (1973). Very similar to C. westi. Biology and sexuales unknown.

Cinara rubicunda (Wilson) Apterae light brown or pinkish, covered with

white powder except for thin spinal stripe (Wilson, 1915); BL c. 3.5-3.6 mm.
On twigs of Juniperus occidentalis in Oregon, USA. Sexuales and life cycle
unknown. Bradley's (1951) account was based on a misidentification of
C. juniperi (Voegtlin and Bridges, 1988).

Cinara russellae Pepper and Tissot Apterae rather shiny black; BL 2.0-
2.5 mm. Found in July in small groups at bases of needles of Pinus banksiana,
feeding on the sheath area rather than on the branch, but in autumn oviparae
and apterae were found in dense colonies with C. nigra on the trunk and
branches (Pepper and Tissot, 1973). Only known from two collections in
Michigan, USA. Oviparae and alate males in late September.

Cinara rustica Hottes Apterae mostly black dusted to grey with wax
powder, except for the posterior abdomen which is dull chocolate brown
(Hottes, 1960b); BL 3.2-3.4 mm. On bark of older twigs or on trunks of Pinus
edulis, with a preference for young vigorous trees. In Arizona and Colorado,
USA. Oviparae (Hottes, 1956d) and apterous males (Hottes, 1960b) in
October.

Cinara saccharinipini Hottes Appearance in life not recorded: BL 3.4-

4.0 mm. On Pinus lambertiana in California, USA. Apterous and alate vivi-
parae collected in December, indicating anholocycly (Hottes, 1958g).
Biometrie data given by Voegtlin (1976) indicate a close relationship to
C. moketa, except for a greater number of accessory hairs on R IV (14-16,
as opposed to 8-9 in moketa).

Cinara saraswatae Das and Raychaudhuri Apterae long-bodied, dark brown;

BL 2.8-3.3 mm. Feeding at bases of needles on tender stems of an unidentified
Pinus sp. in Nepal (B.C. Das and Raychaudhuri, 1983). No other morphs
known.

Cinara saskensis Bradley Apterae dark grey, without wax; BL 5.1-5.6 mm.
On bark of main roots of Picea glauca, near the trunk and just below ground
level (Bradley, 1962). In Manitoba and Saskatchewan, Canada. Life cycle and
sexuales unknown.

Cinara schimitscheki Börner Apterae and alatae broadly oval and somewhat
flattened dorsoventrally, dark brown covered with wax powder; BL 3.3-
5.1 mm. In spring at twig apices among previous year's needles, later on older
branches under bark (Pintera, 1966). Host is usually Pinus nigra (incl, var.
maritima), sometimes P. mugo, P. pinea or P. leucodermis. It occurs through-
out Europe (except Scandinavia and the Baltic), east to Crimea and Turkey.
Life cycle incompletely known; Scheurer (1976) found and described oviparae
 Cinara 647

(and 'sexuparae') in Germany. Binazzi (1978) gave an account of schimitscheki

in Italy, and Michelena Saval and Gonzalez Funes (1988a) recorded its
parasitoids in Spain. 2n = 10*.

Cinara schuhi Hottes Apterae probably with light brown abdomen but
darker head, thorax and legs (Hottes, 1957a); BL 2.5-2.9 mm. On Abies con-
color in Oregon, USA. Not collected since the original description; biology
and sexuales unknown.

Cinara schwarzii (Wilson) Apterae cinnamon-brown, with a black area

between the siphuncular cones, sometimes extending further over abdominal
dorsum, and white spots on thorax and in a spinal row on abdomen (Voegtlin,
1976); BL 2.2-4.2 mm. In medium to large colonies, mostly on one- to three-
year-old branches of Pinus ponderosa and its varieties, also on P. jeffreyi and
P. cooperi, in western USA and Mexico (Ortiz, 1982). Records from other
hosts and localities are suspect. Oviparae and males in September-October;
eggs are laid in rows on the needles (Palmer, 1926; and Fig. 7).

Cinara setulosa Hottes and Essig Appearance of aptera in life unknown,

probably rather pale (males have a bright green abdomen - Hottes, 1957b);
BL of aptera 4.5-5.0 mm. On lower branches of Abies magnifica var.
shastensis in Oregon, USA. Oviparae and alate males in late September.

Cinara shinjii Inouye Apterae have pale brown head and thorax and dark
greyish-brown abdomen, with an extensive pattern of white wax powder
(Moritsu, 1983); BL 2.5-3.0 mm. On twigs, branches and stem of Pinus
parviflora and P. himekomatsu in Japan, forming large colonies on lower
parts of tree in summer (Inouye, 1970). Alatae are produced in July, oviparae
and alate males in September-October. Paik's (1965) account of C. shinjii on
P. koraiensis in Korea is of a different, shorter-haired species.

Cinara sitchensis Hottes Appearance in life unknown; BL of aptera 2.4-

2.6 mm. Other morphs unknown. Described from Picea sitchensis in Califor-
nia, USA (Hottes, 1958b), and also collected in Oregon (BMNH colln).

Cinara (Cupressobium) smolandiae Danielsson and Carter Apterae dusky

brownish-grey with dull bronze metallic highlights in sunlight, and shiny black
siphuncular cones; BL 3.3-4.4 mm. Feeding in bark crevices or rust cankers
1-2 m above ground on stems of old (50-year plus) trees of Juniperus com-
munis. In Sweden (Danielsson and Carter, 1993a). Sexuales and life cycle
unknown.

Cinara solitaria (Gillette and Palmer) Apterae tick-like in behaviour and

appearance, dull brown to yellow-brown, slightly wax-dusted, with six longi-
tudinal rows of black dots (Palmer, 1952); BL 2.5-3.0 mm. On Pinus
ponderosa, living singly on bark of tender shoots, positioned head downwards
at bases of needles. In Arizona (BMNH colln), Colorado and Utah, USA.
Oviparae in early October, males not recorded.
648 Cinara

Cinara sonata Hottes Apterae broadly oval, dull reddish-brown to dark grey
with numerous small irregularly distributed black spots, and at least in the
ovipara there are patches of white wax powder on pronotum and narrow cross-
bands of wax on abdominal tergites (Hottes, 1955c, 1957b); BL of aptera
5.3-7.0 mm. On undersides of small branches of Abies magnifica var.
shastensis, and also recorded from A. concolor, in western North America.
Oviparae and alate males in late September.

Cinara soplada Hottes Appearance in life unknown; BL of aptera c. 3.1 mm.

Apterae collected on an unidentified scrub spruce (Hottes, 1956c), but the
alate male was subsequently described from Picea glauca (Hottes, 1958a).
Only known from Maine, USA. Alate viviparae and oviparae not recorded.

Cinara sorini Inouye Apterae long-bodied, brown to dark brown: BL

6.1-6.8 mm. On Pinus thunbergii in Japan, living on bark of trunk of old
trees, usually close to the ground in ant shelters (Inouye, 1970). Life cycle
and sexuales unknown. Closely related to, and perhaps synonymous with,
C. etsuhoe.

Cinara spiculosa Bradley Apterae dark brown; BL c. 3.0 mm. On small

twigs of Larix laricina, often in loose colonies on lower branches, closely
resembling old buds (Bradley, 1961). Oviparae and alate males in autumn
(Bradley). In Canada, from Alberta across to Nova Scotia.

Cinara splendens (Gillette and Palmer) Apterae rusty-brown, slightly dusted

with wax on head, lateral areas and intersegmentally, with conspicuous black
patches on metanotum and ABD TERG 1 and a pale median dorsal area on
anterior abdomen; BL 2.2-2.6 mm (Gillette and Palmer, 1924). On bark of
twigs of Pseudotsuga menziesii in Colorado, USA. Oviparae and alate males
in September-November. Very similar to C. pseudotsugae, and possibly a
short-haired form of that species.

Cinara strobi (Fitch) Apterae long-bodied, shiny metallic to dull grey-black,

with a spinal stripe and spots of white wax (Pepper and Tissot, 1973); BL
3.0-3.4 mm. Often in large colonies on branches and upper trunk of Pinus
strobus throughout its range in eastern North America. Records from P. bank-
siana and P. resinosa need confirmation (MacGillivray, 1955). Oviparae and
alate males in September-November. Eggs are laid in rows on needles (Bradley,
1951). Weigel and Baumhofer (1948) described damage to white pines.
2n = 10*.

Cinara subapicula Zhang Appearance in life unknown; BL of aptera

c. 3.3 mm. A long-haired species described (alatae only) from unidentified
Pinus spp. in Tibet (Zhang and Zhong, 1981b).

Cinara subterranea Bradley Only the sexuales are recorded (Bradley, 1956a).
Oviparae are greyish-brown with dark brown legs and antennae, BL c. 3.3 mm,
 Cinara 649

collected mid-September. On roots of Larix laricina in Alberta and Manitoba,

Canada, forming dense colonies (Bradley, 1961). Males are alate.

Cinara taedae Tissot Apterae dark brown to brownish-black, often shining;

BL 2.1-3.1 mm. Feeds on bark between needle bases of one- or two-year-old
growth, or sometimes on current terminal shoots (Pepper and Tissot, 1973).
On Pinus spp. of subsections Australes and Contortae, especially P. taeda and
P. rigida, in western USA south from Pennsylvania. Oviparae and alate males
on P. rigida in Pennsylvania in October-November.

Cinara taiwana (Takahashi) Apterae blackish-brown, shining; BL c. 3.8 mm.

On Tsuga formosana at a high altitude in Taiwan (Takahashi, 1925). Biology
and sexuales unknown.

Cinara tanneri (Knowlton) Appearance in life not recorded; BL of aptera

3.3-4.0 mm. On pinyon pines (P. edulis, P. monophylla) in Utah, USA.
Sexuales and biology unknown and the alatae in the original description were
C. edulis according to Hottes (1960b).

Cinara tenuipes Chakrabarti and Ghosh Appearance in life not recorded;

BL of aptera 3.6-4.3 mm. Originally described as a subspecies of abieticola
( = confinis), from a host plant uncertainly identified as Abies (Chakrabarti
et al., 1974) in Himachal Pradesh, India. However, Starý and Raychaudhuri
(1982) described two new species of parasitoid (Pauesia) from this aphid and
the host is given as Pinus sp. Paratypes in the BMNH collection seem to be
small, long-haired specimens of the eastopi/orientalis group.

Cinara terminalis (Gillette and Palmer) Apterae light cinnamon-brown,

sometimes tinged with green, darkest in centre of dorsum, with some wax
dusting on head, thorax and intersegmentally on abdomen (Gillette and
Palmer, 1924); BL 2.2-2.4 mm. Described from specimens living solitarily and
inconspicuously on bark of young tender twig terminals of Pinus edulis in
Colorado, USA. Also recorded from Utah, Arizona, Nevada and Mexico. The
Mexican records (Ortiz, 1982) are from Pinus cembroides and several other
Pinus spp., including michoacana and pseudostrobus in subsect. Ponderosae.
Paratypes of C. terminalis in BMNH collection resemble small C. ponderosae,
and the distinction between these two species needs to be clarified.

Cinara thatcheri Knowlton and Smith Appearance in life unknown; BL of

aptera 3.2-4.5 mm (Knowlton and Smith, 1938). On Pinus ponderosa in
western USA and British Columbia. Biology and sexuales unknown. Very
similar to, and possibly synonymous with, C. schwartzii.

Cinara tibetapini Zhang Appearance in life unknown; BL of aptera

c. 6.5mm. On Pinus sp. in Tibet (Zhang and Zhong, 1981b). Other morphs
and biology unknown.
650 Cinara

Cinara tistaensis Agarwala and Raychaudhuri Appearance in life unknown;

BL of aptera 3.7-4.0 mm. On an unidentified Abies sp. in Sikkim, India
(Agarwala and D. Raychaudhuri, 1982). Other morphs and biology unknown.

Cinara todocola (Inouye) Apterae have brown head and dark green to
greenish-black abdomen, lightly dusted with wax (Inouye, 1936); BL 2.6-
3.0mm. On trunk and branches of young todo-fir trees (Abies spp. in Japan
and Sakhalin). Often in ant shelters. Injurious. Oviparae and alate males in
September-November (Inouye, 1970). Yamaguchi (1976) studied ecology and
morph determination, and Furuta (1984) studied the impact of natural
enemies. Cinara smaragdina is a similar species on Abies spp. in Siberia, but
apparently with a different biology as it feeds on shoots between needle bases
(Pashchenko, 1988b).

Cinara tonaluca Hottes and Wehrle Apterae dusted with white wax, with a
black stripe and two longitudinal rows of black spots showing through the wax
(Hottes, 1952b); BL 2.1-2.6 mm. On woody stems of young trees of Juniperus
monosperma in Arizona, USA. Not recorded since the original description
(Hottes and Wehrle, 1951), sexuales and life cycle unknown.

Cinara tsugae Bradley Apterae have abdomen chestnut-brown, head and

thorax somewhat lighter brown; BL 2.9-3.6mm. In dense spring colonies on
branches of Tsuga heterophylla and later on roots or lower part of trunk in
ant shelters (Bradley, 1960). In British Columbia, Canada and Oregon, USA
(BMNH colln, leg. Bradley). Oviparae deposit eggs on needles in autumn,
males are unrecorded. Cinara wahsugae, described from Tsuga mertensiana
in Oregon (Hottes, 1960a), may be a synonym, although oviparae and alate
males of wahsugae were collected in August (and see Hottes, 1961c).

Cinara (Cupressobium) tujafilina (Del Guercio) (= winonkae Hottes)

Apterae reddish-brown with a dorsal pattern of bluish-white wax and two dark
brown divergent curved bands running from head to about level of siphunculi;
BL 1.7-3.5 mm. On Cupressaceae; Callitris, Chamaecyparis, Cupressus, Juni-
perus (but not communis), Thuja (usually orientalis) and Widdringtonia. On
foliated branches, undersides of branches near the trunk, on wound tissue,
or in midsummer on roots (Bray, 1953; Colombo and Parisini, 1984). Virtually
cosmopolitan, including many warmer regions. Apparently entirely anholo-
cyclic; sexuales are not recorded. There are recent population studies by
Furuta (1988; Japan) and Mohammed et al. (1988; Iraq). Similar but shorter-
haired specimens in southwest Asia may be referable to C. mediterraneum
Narzikulov. 2n = 12.

Cinara vagabunda Hottes and Essig Appearance in life unknown; BL of

aptera c. 3.6mm. On Pseudotsuga menziesii in Arizona and New Mexico,
USA (Hottes, 196Id). See C. pseudotaxifoliae.

Cinara vandykei (Wilson) Apterae dark brown to black with little or no

wax; BL 1.8-3.0 mm. In colonies on bark of twigs of Picea spp. in western
 Cinara 651

USA (incl. California and Oregon: BMNH colln) and across Canada. Ovi-
parae and apterous males are produced early, in July-August (Palmer, 1926).
Cinara wanepae Hottes, described from Picea pungens in Colorado, is a pro-
bable synonym (Palmer, 1936b).

Cinara villosa Gillette and Palmer Apterae described as indistinguishable in

life from those of C. apini (Gillette and Palmer, 1924) and therefore pre-
sumably dark brown, mottled with wax powder; BL c. 3.2-4.0 mm. On bark
of twigs of Pinus flexilis in western North America. Originally described as
a long-haired form of apini in Colorado and possibly the two are not speci-
fically distinct. Hottes (1955b) described two variant populations of C. villosa
from P. flexilis var. reflexa in Arizona as subspecies.

Cinara wahhaka Hottes Apterae shining dark brown without wax; BL 1.5-
1.8mm. On foliage of small twigs of Juniperus osteosperma ( = utahensis),
only known from Colorado, USA. All the second generation were alate
(Hottes, 1952c). Oviparae and apterous males (Hottes, 1953b) in September-
October.

Cinara wahluca Hottes Apterae reddish-brown without wax; BL 2.7-

3.0mm. On bark of trunk and larger branches of Juniperus scopulorum,
under scales or in crevices or wounds. Only known from Colorado, USA.
Oviparae and apterous males in September (Hottes, 1952a).

Cinara wahsugae Hottes See C. tsugae.

Cinara wahtolca Hottes Apterae brownish to dark grey with black marking
and black siphunculi, usually extensively covered with white powder (Hottes,
1953a); BL 2.6-4.6 mm. In large colonies on trunk and needle-free limbs of
Pinus spp. of subsect. Cembroides (pinyon pines) in southwestern USA.
Oviparae and alate males in October (Hottes, 1954b, 1956e). Hottes (1955b)
described a short-haired form on P. edulis in Arizona as a subspecies,
C. wahtolca curtiwahtolca; similar short-haired specimens have also been
collected in Utah on P. monophylla (BMNH colln).

Cinara watanabei Inouye Apterae have head and thorax dark brown,
abdomen shiny brown; BL 4.8-5.4 mm. In large colonies on three- to five-
year-old wood of branches of older trees, or on trunk of young trees, of Pinus
spp. (e.g. koraiensis, parviflora) in Japan, Korea and Sakhalin (Pashchenko,
1988b). Oviparae and alate males in Japan in October-November (Inouye,
1970). Cinara pinikoraiensis, described from China (Zhang and Zhong, 1989),
is possibly a synonym.

Cinara watsoni Tissot Apterae have head and thorax dark chestnut-brown,
abdomen cinnamon-brown with dark brown markings, the whole body dusted
with greyish-white wax (Tissot, 1939); BL 2.7-4.3 mm. In small loose groups
on new shoots, or near ends of previous year's growth, of Pinus spp. of
652 Clethrobius

subsect. Australes in eastern USA. Oviparae and alate males occur in October-
November (Pepper and Tissot, 1973). Population trends of C. watsoni in
South Carolina were studied by Brooks and Warren (1964), and by Patti and
Fox (1981).

Cinara westi Tissot and Pepper Apterae dark brown; BL 2.1-2.8 mm. On
bark of Pinus taeda in southwestern USA, often associated with lesions of
the rust Cronartium fusiforme and usually in ant (Crematogaster) shelters
(Tissot and Pepper, 1967). Sexuales and life cycle unknown. This species is
possibly synonymous with C. rigidae.

Cinara zoarcbursara Knowlton Appearance in life unknown; BL of aptera

c. 3.5mm. Only known from apterae collected on Abies concolor in Utah,
USA (Knowlton, 1935). Biology unknown.

CLETHROBIUS Mordvilko Drepanosiphinae: Phyllaphidini

Three or four species of large (2.8-5.6 mm) hairy brown aphids living on twigs
and young branches of Alnus and Betula species. All adult viviparae and males
are alate, the ovipara being the only apterous morph. Accounts are available
for Europe (Stroyan, 1977; Heie, 1982; Wood-Baker, 1983) and India (Chak-
rabarti, 1988).

Clethrobius comes Walker Alate viviparae are dark brown, forming clusters
on branches and twigs of Betula spp., often where the new growth is dying
back, or on twigs of Alnus spp. overhanging streams. The populations on
Alnus are regarded by some authors as a separate species, C. giganteus
Cholodkovsky, and attempts to transfer aphids from birch to alder have not
succeeded (Wood-Baker, 1983). No consistent morphological differences have
been found, however, and both birch and alder populations share the same
peculiar structurally heterozygous karyotype (2n = 11; Blackman, 1988).
Clethrobius comes occurs throughout Europe and across Asia to Korea and
Japan. Holocyclic, with sexuales in October-November.

Clethrobius dryobius Chakrabarti and Raychaudhuri Brown aphids with

many more hairs on the dorsal surface of the body and the last rostral
segments than C. comes, and also with some hairs on the siphunculi.
Holocyclic on Betula alnoides and B. utilis in northern India, with sexuales
in October-November (Chakrabarti, 1988; Chakrabarti et al., 1988).

Clethrobius vermai Ghosh and Quednau Colour in life not recorded.

Described from a single alata from Betula sp. in Himachal Pradesh, India
(A.K. Ghosh and Quednau, 1990).
 Colopha 653

CLYDESMITHIA Danielsson Pemphiginae: Pemphigini

A genus with one nearctic species related to Pachypappa or Epipemphigus.

The fundatrices lack wax glands and have antennae with only 4 segments.

Clydesmithia canadensis Danielsson Gall on Populus (balsamifera, tricho-

carpa) consists of an elongate slightly wrinkled swelling on the distal half of
the leaf lamina, 1.0-1.5 cm long, tinged with red, opening on underside of
leaf. Apterae in galls are yellowish with a thin layer of wax; BL 1.5-2.7 mm.
In Canada (Alberta, British Columbia). Apparently monoecious on Populus;
alatae and sexuales unknown (Danielsson, 1990c).

COLOPHA Monell Pemphiginae: Eriosomatini

A genus related to Kaltenbachiella. Eastop and Hille Ris Lambers (1976) listed
eight species, but after revisions by Akimoto (1985a) and Smith (1985) only
four species now remain in Colopha, three host-alternating between Ulmus
and Cyperaceae (see below), and one (kansugei) anholocyclic on Carex. One
additional species, caucasica, is included here although it differs biologically
and its true generic position is uncertain.

Colopha caucasica (Dzhidlabze) Spring generations roll and blister the

leaves of Zelkova acuminata and Z. carpinifolia (Dzhibladze, 1960). Alatae
(BL 1.3-1.8 mm) emerge from the galls in May and fly to roots of Carex
(BMNH colln, leg. D. Hille Ris Lambers), on which sexuparae are produced
in November. In southwest and Central Asia (Georgia, Iran; BMNH colln).

Colopha compressa (Koch) Forming pouch galls on upper surfaces of leaves

of Ulmus spp., near the mid-rib (Fig. 134B, p. 898); they are laterally com-
pressed, cockscomb-shaped, yellowish, often tinged with red. Alatae (BL
1.4-1.8 mm) emerge in July and colonize roots of Carex and Eriophorum.
Sexuparae return to elm in September-October and anholocyclic popula-
tions may also persist on roots of Cyperaceae through the winter months
(Marchal, 1933; Zwölfer, 1957). Throughout Europe (incl. Iceland), east
to Turkey and Ukraine and introduced into towns in Siberia (Heie, 1980).
2n = 16.

Colopha graminis (Monell) Forming elongate cockscomb-like galls on upper

surfaces of leaves of Ulmus spp. (americana, fulva) on the leaf lamina between
654 Colophina

the veins (Fig. 134C, p. 898). Alatae (BL 1.0-1.6 mm) emerge in June-July,
migrating to roots of Leersia spp.; an attempted transfer to Eragrostis was
unsuccessful (Patch, 1910b, cf. ulmicola). Secondary hosts must also include
Aira caespitosa and Agrostis plumosa, from which the species was originally
described (Monell, 1882). Sexuparae develop to adult on the leaves of the
secondary host, returning to elm in October (Patch, 1910b). Widely distributed
in North America. Some authors (e.g. Palmer, 1952) have synonymized
graminis with ulmicola, but Smith (1985) considered that there was sufficient
evidence to regard the two species as distinct.

Colopha ulmicola (Fitch) Forming elongate cockscomb-like galls indistin-

guishable from those of C. graminis on leaves of Ulmus spp. (americana,
fulva, thomasii). Alatae (BL 1.0-1.6 mm) emerge in June-July and fly to
Eragrostis spp., giving rise to colonies with flocculent wax on the upper
leaves and stems. Possibly other Gramineae are also colonized (but not
Leersia! - see Patch, 1910b). Sexuparae return to elm in September-November
(Patch, 1910b). Widely distributed in North America.

COLOPHINA Börner Pemphiginae: Eriosomatini

Four East Asian species related to and morphologically resembling Eriosoma

but biologically distinct (Akimoto, 1983), with particularly complex life cycles
and polymorphism (Aoki, 1980b). Two species are known to alternate between
Zelkova and the aerial parts of Clematis, while the other two are only known
from Clematis.

Colophina arma Aoki The galls are globular, loosely closed, on the margins
of leaves of Zelkova serrata in Japan (Fig. 125B). Alatae (BL 2.2-2.4 mm)
emerge in July and fly to Clematis stans. where dense colonies (including
sterile 1st instar soldiers) are formed on stems or peduncles. Sexuparae are
produced in autumn and presumably return to Zelkova, but overwintering also
occurs as midget first instar larvae under bark of the secondary host (Aoki,
1980b). 2n (female) = 10, 2n (male) = 8 (Blackman, 1986).

Colophina clematis (Shinji) The galls on Zelkova serrata in Japan appear

identical to those of C. arma (see above). Alatae (BL c. 2.2 m) emerge in July
and fly to Clematis apiifolia, where dense colonies (including soldiers) are
formed on the stems, usually in shade near the ground (Aoki, 1977). Alate
sexuparae return to Z. serrata in October, but 1st instar larvae also overwinter
in bark crevices on the secondary host (Aoki, 1980b). 2n (female) = 11, 2n
(male) = 9 (Blackman, 1986).
 Cornaphis 655

Fig. 125. Galls of Eriosomatini on Zelkova. A, Byrsocryptoides zelkovaecola on Z. carpinifolia (after

Dzhibladze, 1965b); B, Colophina arma on Z serrafa (viewed from underside, after Aoki, 1980b);
C, Paracolopha morrisoni on Z serrata (after Akimoto, 1985a).

CORNAPHIS Gillette Pemphiginae: Pemphigini

A genus for one North American species monoecious on Populus, charac-

terized by a medial frontal projection that forms a distinct horn in apterae
and a raised base for the median ocellus in alatae. (Cornaphis viscisucta
Zhang, described from Viscum album in China (Zhang and Zhong, 1985b) is
unlikely to belong to this genus.)

Cornaphis populi Gillette Galls on Populus angustifolia are crescent-shaped

folds of the leaf-edge, pale or streaked with red. In western USA (Wyoming,
Colorado, Utah). Apterae inhabit the same galls as the slate-grey fundatrices
and are straw-yellow with black head and four dark patches on the pronotum,
dusted with wax. There is no host alternation; alate sexuparae (BL
2.0-2.3 mm) are produced in the galls in July (Gillette, 1913; Palmer, 1952).
656 Corylobium

CORYLOBIUM Mordvilko Aphidinae: Macrosiphini

A genus for one Macrosiphum-like species with capitate hairs living on

Corylus. Hille Ris Lambers (1947a) discusses its affinities.
Corylobium avellanae (Schrank) Apterae green, sometimes reddish, spindle-
shaped; BL 1.7-2.7 mm. On shoot apices, petioles and undersides of young
leaves of Corylus avellana; dense colonies form particularly on suckers. In
Europe eastward to Turkey and Ukraine and introduced into Canada (British
Columbia), where it was also found on C. cornuta and C. maxima (Forbes
and Chan, 1984). Monoecious holocyclic, with alate males. Natural enemies,
especially the impact of a pathogen, were discussed by Viggiani in Cavallaro
(1983) and Tremblay and Pennachio (1985) reported a parasitoid (Praon dor-
sale). See also Hille Ris Lambers (1947a). 2n = 10.

CRANAPHIS Takahashi Drepanosiphinae: Phyllaphidini

A little-known oriental genus of one or two species related to Takecallis and

associated with Arundinaria. Shivaphis arundinariae Takahashi was trans-
ferred to Cranaphis by Tao (1958), but returned to Shivaphis by Liao (1976).
Cranaphis formosana (Takahashi) All viviparae alate, long-bodied, yellow
with black antennae and a broad medial dusky stripe on head and prothorax
and paired dusky markings on most abdominal tergites (Takahashi, 1924a);
BL 1.5-2.2mm. On undersides of leaves of Arundinaria (= Yushania)
niitakayamensis in the central mountains of Taiwan (Liao, 1976). Sexuales
unknown; probably anholocyclic in Taiwan. Cranaphis indica Chakrabarti
and Raychaudhuri (1976), described from a single alata and immatures
collected on Arundinaria sp. in West Bengal, India, is very similar to C.
formosana, and may be a synonym.

CRYPTURAPHIS Silvestri Drepanosiphinae: Phyllaphidini

Alnus-feeding aphids with peculiar projections of the head and prothorax.

Crypturaphis grassii Silvestri Monoecious holocyclic on Alnus cordata
in Italy and Corsica. Dorsoventrally flattened, brownish adult apterae and
 Dermaphis 657

green immatures are dispersed along veins on both upper and lower leaf
surfaces. Alatae, the most commonly collected morph in summer, have black
head and thoracic lobes, paler prothorax and an ill-defined and variably
developed brown patch across ABD TERG 5-6; BL 2.2-3.0 mm. Oviparae
and males occur in October-November. Patti (1983) gives some biological
information.

DASYAPHIS Takahashi Drepanosiphinae: Phyllaphidini

A genus for one oriental species associated with Juglandaceae, the apterae of
which have fused head and prothorax, reduced antennae and long hair-bearing
processes - although, according to Tseng and Tao (1938), an apterous morph
without such tubercles can also occur in colonies.

Dasyaphis rhusae (Shinji) Apterae are dorsoventrally flattened, pale

yellowish-green, yellow or white; BL 1.2-1.4 mm. On undersides of mature
leaves of Juglans spp. (ailanthifolia, mandshurica) in Japan (Takahashi,
1965b; Higuchi, 1972; as Dasyaphis onigurumi), Korea (Paik, 1972), Siberia
(Quednau and Shaposhnikov, 1988) and also recorded from Pterocarya
stenoptera in China (Tseng and Tao, 1938; as Sinocallis mirabilis).

DERMAPHIS Takahashi Hormaphidinae: Nipponaphidini

Species related to Thoracaphis, only known from the apterous viviparae on

Fagaceae, which have a heavily sclerotized and sculptured dorsal cuticle and
long pointed marginal prosomal hairs. Siphunculi are lacking and the tarsi are
often without claws. Descriptions and available specimens are inadequate to
construct a key to species. We follow A.K. Ghosh and Raychaudhuri (1973a)
in including three species in this genus, although this is mainly for convenience
as these species do not seem to be very closely related.

Dermaphis crematogastri (Takahashi) Apterae black, with some wax. Body

oval, convex dorsally, extended somewhat at posterior end, sometimes asym-
metrical and irregular in outline; BL c. 1.0-1.1 mm. The prosoma has many
long stout mediodorsal as well as marginal hairs. Tarsi usually have claws. On
branches of Quercus sp. in Thailand, attended by ants (Crematogaster sp.),
and often in their shelters (Takahashi, 1941; as Thoracaphis).

Dermaphis japonensis Takahashi Apterae black, with some greyish-white

wax. Body oval, convex dorsally, extended slightly at posterior end; BL
658 Dinipponaphis

1.1-1.8 mm. Even the hind legs lack claws and are often concealed beneath
body. Dorsal median area of prosoma with only 4 pairs of hairs, which are
shorter than the marginal hairs (Takahashi, 1958a). On branches of Quercus
glauca and Lithocarpus sp. (Shinji, 1941; as Thoracaphis takahashii) in Japan
and on Quercus sp. in Korea (Paik, 1972). Ant attendance is not recorded.

Dermaphis takahashii (Strand) Apterae black, with white wax around

margin. Body subcircular, deeply convex; BL c. 1.2mm. The hind legs
project behind the body and the tarsi always have claws. In small groups
on branches and stems of Quercus glauca, Quercus sp. and Lithocarpus
sp., usually in ant shelters made by Crematogaster, in Taiwan and China
(Takahashi, 1935d, as Thoracaphis; Tao, 1966).

DINIPPONAPHIS Takahashi Hormaphidinae: Nipponaphidini

A genus for one species related to Nipponaphis, living without host alternation
on Distylium. The alate sexuparae have clear wings and characteristic sensoria-
tion of the antennae. A.K. Ghosh and Raychaudhuri (1973a) revised the genus.

Dinipponaphis autumna (Monzen) In small reddish galls, projecting on

both sides of leaves, of Distylium racemosum in Japan (Fig. 130A, p. 785; and
Moritsu, 1983). Probably also in Korea, as some of the galls illustrated as
Neothoracaphis yanonis by Paik (1972; his Figs262a, b) seem to be this
species. Monoecious holocyclic. Alate sexuparae emerging from the galls in
early December have black head and thorax, clear wings with pale veins and
pterostigma and yellow abdomen; BL 1.5-1.7 mm (Takahashi, 1962a). They
deposit males and oviparae of very different sizes (Sorin, 1987b, 1990a) on
the undersides of the leaves. Adult sexuales overwinter on the buds, eggs are
laid in March and fundatrices hatch in April (Sorin, 1960).

DIPHYLLAPHIS Takahashi Drepanosiphinae: Phyllaphidini

A genus of four palaearctic and one nearctic oak-feeding species, related to

the wholly nearctic genus Stegophylla, but the siphunculi are small pores
without encircling hairs. Higuchi (1972) revised the Japanese species and
Quednau (1971) keyed all five species.

Diphyllaphis alba Takahashi Appearance in life not recorded, presumably

entirely white with flocculent wax; BL 1.1-1.4 mm. On Quercus phyllyraeoides
in Japan (Honshu, Kyushu; Takahashi, 1960c; Higuchi, 1972). Only the
 Distylaphis 659

apterous viviparae are known, collected in April (BMNH colln, leg. R. van
den Bosch) and September-October.

Diphyllaphis konarae (Shinji) Apterae elongate pear-shaped, light red,

orange or white, with white flocculent wax; BL 1.2-1.7 mm. On Quercus spp.
in Japan (Honshu), on rather mature leaves that curl up and may be bound
together by the wax secreted by the aphid (Shinji, 1924; Takahashi, 1960c).
Only apterous viviparae are known, collected in October.

Diphyllaphis microtrema Quednau Apterae are broadly oval, pale yellowish-

green, with dense white flocculent wax; BL 1.7-2.2 mm. In dense woolly masses
on undersides of leaves of Quercus rubra in eastern USA (Connecticut, North
Carolina). Holocyclic; oviparae and alate males in September (Quednau, 1971).

Diphyllaphis mordvilkoi (Aizenburg) Apterae are elongate pear-shaped,

yellowish-white, with much flocculent white wax; BL 1.2-1.5 mm. On under-
sides of mature leaves of Quercus spp., causing yellowing and necrosis
(Binazzi and Roversi, 1988). In Spain, Italy (Tuscany, Sicily), Turkey,
Lebanon (BMNH colln) and Caucasus. Holocyclic, with apterous males
(Roversi and Binazzi, 1990).

Diphyllaphis quercus (Takahashi) Apterae are elongate pear-shaped, whitish

with flocculent wax; BL 1.4-1.6mm. On Quercus spp., curling the tips of
the leaves and causing necrosis. In Japan, China and Korea. Moritsu (1983)
described and illustrated the alate viviparous female. Oviparae occur in Korea
in early November (Paik, 1972, as Nymphaphis quercus).

DISTYLAPHIS Noordam Hormaphidinae: Nipponaphidini

One species producing galls on Distylium in Java, but also having free-living
generations on the same host. First instar 'soldiers' occur in the galls
(Noordam, 1991).

Distylaphis foliorum (van der Goot) Galls on Distylium have not been
described. Alatae emerging from galls in late January have BL 2.6-3.3 mm,
long forewings with a broad black pterostigma and the wing membrane
fuscous basad of Cu lb , and secondary rhinaria distributed III 20-26, IV
8-12, V 7-10. Apterae are bright greenish, margins of body with thick
powdery or flocculent wax, and with 2 longitudinal rows of wax patches or
a thick coat of wax on dorsum; BL c. 1.5mm. On undersides of leaves of
Distylium stellare in Java. Apparently monoecious holocyclic, but life cycle
needs further clarification. Oviparae are present on Distylium leaves in
April-August; males are unrecorded (Noordam, 1991).
660 Doraphis

DORAPHIS Matsumura and Hori Hormaphidinae: Cerataphidini

A genus for one or two poplar-feeding species with no obvious close relatives,
the large aleyrodiform apterae of which were originally described as a scale
insect. Alatae have an unbranched media in the forewing and a single oblique
vein in the hindwing. Tao (1969) gave an account of the genus (in Chinese).

Doraphis populi (Maskell) Apterae dark, flattened, scale-like, almost cir-

cular, secreting wax from a ring of marginal glands; BL 2.2-3.0 mm. On bark
of Populus spp. (incl. ciliata, maximowiczii, sieboldii). Alatae are purplish-
brown, with rather short 4- or 5-segmented and heavily sensoriated antennae;
BL 2.2-2.6 mm (Matsumura and Hori in Hori, 1929). Immature alatae form
groups on undersides of leaves (Takahashi, 193la). In Japan, China, Korea,
Siberia and India. Anholocyclic populations on P. tremulae in the region of
Vladivostock have only two parthenogenetic generations a year, one apterous
and one alate (Mordvilko, 1929a, as Paracerataphis tremulae; made a sub-
species of populi by Matsumura and Hori in Hori, 1929). Elsewhere the life
cycle is unknown. Ceratoglyphina populisucta Zhang and Zhong (1985d) is
possibly a previously undescribed morph of this species.

DREPANAPHIS Del Guercio Drepanosiphinae: Drepanosiphini

Active, rather robust alate aphids on leaves of nearctic Aceraceae. Apparently

related to the oriental genus Yamatocallis and with similar flask-shaped
siphunculi, which differ however in the absence of any subapical reticulate
ornamentation. Dorsal abdominal tubercles are variably developed on ABD
TERG 1-4 (usually largest on 3) and these are often conspicuously pigmented.
Holocyclic, with males and all viviparous females alate, oviparae apterous.
Trioxys ameracis is a specialized parasitoid. Smith and Dillery (1968) revised
the genus in a very accessible publication, so only a summary treatment is
provided here, with emphasis on the recognition of the living aphids.

Drepanaphis acerifoliae (Thomas), Plate 7b Alatae have head and thorax

reddish-brown, abdomen pale with dorsal tubercles dark at least on ABD
TERG 3-4, dark siphunculi, and wing veins distinctly dark-bordered. There
are 3 longitudinal white wax stripes on head and pronotum, white wax
markings on meso- and metathorax and anterior abdomen, and a covering
of wax on abdomen posterior to siphunculi. BL 1.9-2.3 mm. On Acer sac-
charinum and A. rubrum, occasionally on A. saccharum and other maples.
Common and widely distributed in North America. 2n = 38.
 Drepanaphis 661

Drepanaphis carolinensis Smith Alatae similar to acerifoliae in life, but the

wing veins are not dark-bordered; BL 1.6-2.2 mm. On Acer saccharum, and
occasionally on A. rubrum. Widespread in North America east of Rocky
Mountains.

Drepanaphis choanotricha Smith and Dillery Alatae black with pale legs.
Head and pronotum striped with white wax, meso- and metanotum and dorsal
abdomen with wax dots, denser posteriorly; BL 1.1-1.7 mm. On low branches
of Acer saccharum (always within 50 cm of ground), in Illinois and North
Carolina, USA.

Drepanaphis granovskyi Smith and Dillery Alatae almost entirely unpig-

mented, without conspicuous wax; BL 1.2-2.0 mm. On Acer grandidentatum
in Idaho and Utah, USA.

Drepanaphis idahoensis Smith and Dillery Alatae entirely covered with

white wax except for a dark, U-shaped mark between the dorsal tubercles
an ABD TERG 3 and the siphunculi. Legs pale, BL 1.4-2.0 mm. On Acer
grandidentatum in Idaho and Utah, USA.

Drepanaphis kanzensis Smith Alatae yellowish-white, with a profuse cover-

ing of white wax, except on thoracic lobes and the prominent dark tubercles on
ABD TERG 3. Legs pale, wing veins not dark-bordered. BL 1.2-2.0 mm. On
Acer saccharum in North America east of Rocky Mountains, but with a more
northerly range than D. carolinensis.

Drepanaphis keshenae Granovsky Alatae have body profusely covered with

white wax, through which project the large dark tubercles on ABD TERG 3.
Wing veins distinctly bordered with fuscous. BL 2.0-2.5 mm. On Acer
saccharum in eastern USA and Ontario, Canada.

Drepanaphis knowltoni Smith and Dillery Alatae white with wax, very like
idahoensis except for dark legs; BL 1.6-2.4 mm. On Acer grandidentatum in
Idaho and Utah, USA and also recorded from other Acer spp. in eastern USA
(Smith and Dillery, 1968).

Drepanaphis monelli Davis Alatae have body entirely covered with white
wax powder except for dark thoracic lobes and a brownish-yellow U-shaped
line between the tubercles on ABD TERG 3 and the dark siphunculi; BL
1.9-2.3 mm. On Aesculus glabra, especially on yellowed leaves (the only
Drepanaphis species not associated with Acer). In eastern USA and Canada
(New Brunswick).

Drepanaphis nigricans Smith Alatae have black body with 2-3 longitudinal
wax stripes on head and pronotum and small spots or flecks of wax on
thorax and abdomen. Legs pale, wings clear except for dark spots at ends
of veins. On Acer rubrum, especially on terminals of rapidly growing shoots
662 Drepanaphis

(water-shoots), in eastern USA, west to Michigan and Tennessee, and Canada

(New Brunswick). Closely related to, and possibly conspecific with, D. tissoti.

Drepanaphis parva Smith Alatae have reddish-brown head and thorax, the
head and pronotum having 5 longitudinal white wax stripes, and greenish-
grey-brown abdomen with white posterior tergites and longitudinal rows of
white wax dots; BL 1.4-2.6 mm. Legs pale, forewings somewhat cloudy with
veins diffusely bordered. On Acer rubrum in eastern USA and Canada.

Drepanaphis sabrinae Miller Alatae are orange-brown to orange-yellow

with small flecks of white wax, and 2-3 longitudinal stripes on pronotum; BL
2.0-2.6 mm. On Acer saccharum in eastern USA, west to Minnesota and
Kansas.

Drepanaphis saccharini Smith and Dillery Alatae very similar in life to

parva (q.v.), but with clear wings; BL 1.5-2.3 mm. On Acer saccharinum (but
not cultivated varieties) in eastern and mid-western USA; only occasionally
on A. rubrum.

Drepanaphis simpsoni Smith Alatae have reddish-brown head and thorax

with short fine longitudinal white wax stripes, and light yellow to white
abdomen with the finger-like tubercles on ABD TERG 1 reddish-brown; BL
1.5-2.3 mm. On Acer saccharum in eastern USA as far south as North
Carolina, and in eastern Canada. Drepanaphis pallida Richards seems to be
a pale form of D. simpsoni. 2n = 30*.

Drepanaphis spicata Smith Alatae in life closely resemble those of

knowltoni (q.v.), but are larger and have fore femora much paler ventrally
than dorsally; BL 2.4-3.2 mm. On Acer spicatum in eastern USA (moun-
tainous areas west to Tennessee and south to North Carolina) and Canada
(New Brunswick). Records from A. grandidentatum in western USA (e.g.
Palmer, 1952) are probably referable to D. knowltoni and/or D. idahoensis.

Drepanaphis tissoti Smith Alatae black with pale legs, strongly resembling
nigricans (q.v.); BL 1.3-2.0 mm. On Acer rubrum in eastern USA, usually on
growing terminals of young shoots (water-shoots); one record from A. sac-
charum. Doubtfully distinct from D. nigricans.

Drepanaphis utahensis Knowlton and Smith Alatae yellow, with head and
pronotum edged with black, thoracic lobes brown, tubercles on ABD TERG
3 dark and entire body frosted with white wax; BL 1.5-2.0 mm. A very pale
form occurs in midsummer. On Acer grandidentatum in western USA (a
record from A. saccharum in Nova Scotia, Canada, seems likely to be another
species). 2n = 30.
 Drepanosiphum 663

DREPANOSIPHONIELLA Davatchi, Hille Ris Lambers

and Remaudière Drepanosiphinae: Drepanosiphini

A genus for one palaearctic species related to the nearctic genus Drepanaphis,
but the primary rhinaria have no fringe of hairs and dorsal body hairs are long
and stiff as in Chaitophorinae. Both apterous and alate viviparae occur.

Drepanosiphoniella aceris Davatchi, Hille Ris Lambers and Remaudière

Apterae grey to dark brown, lightly dusted with wax powder; BL 1.7-2.5 mm.
Femora are brown conspicuously mottled with darker-brown and siphunculi
are dark brown. The siphunculi of alatae are often much longer than those
of apterae and are usually markedly curved; alatae also have a very short wide
pterostigma with two black patches surrounding a pale central area. On under-
sides of leaves of Acer spp. ((cinerascens, microphyllum) in the Mediterranean
region and southwest Asia (Iran, Iraq, Lebanon, Morocco, Turkey). Life cycle
and sexuales of D. aceris s. str. unknown. Two subspecies have been
described, D. aceris fugans on Acer campestre and A. monspessulanum in
France (Remaudière and Leclant, 1972; with longer antennal hairs, longer
antennal processus terminalis, longer siphunculi and unconstricted CAUDA),
and D. aceris caucasica on Acer ibericum in Armenia (Mamontova, 1982; no
wax powder, straight siphunculi of similar length in both apterae and alatae).
Possibly these are all populations of one very variable species. Drepanosi-
phoniella aceris fugans is holocyclic in France, producing apterous males
and oviparae on A. campestre in October (Remaudière and Leclant, 1972).
Starý and Mackauer (1971) described a new species of parasitoid from this
subspecies.

DREPANOSIPHUM Koch Drepanosiphinae: Drepanosiphini

A genus for eight species of medium-sized to large elongate aphids with long
antennae and tubular siphunculi, all associated with Acer. All viviparae are
alate, forming 'spaced-out aggregations' on undersides of leaves in shade
(Kennedy et al., 1967). Males are alate, oviparae are large and apterous with
the end of the abdomen extended like an ovipositor (Fig. 6). Discritulus
planiceps, which pupate like Praon in a tent under the aphid, Monoctonus
pseudoplatani and Trioxys cirsii (which emerge from pale mummies), and
Aphelinus thomsoni (which emerge from black mummies) are specialized
parasitoids. Stroyan (1977) provided a generic diagnosis and keys to British
species, and Heie (1982) gave an account of the species of northwest Europe.
664 Drepanosiphum

Drepanosiphum acerinum (Walker) Alatae whitish-green or yellow, with

thoracic lobes darker, a conspicuous brown-black spot in front of the base
of each siphunculus, siphunculi either entirely or distally dark, and often a
short brown bar or bars on ABD TERG 4-5 only; BL 2.1-3.3 mm. On Acer
pseudoplatanus, under lower leaves, usually in shade. In Europe, except the
Baltic region. Sexuales in September-October. Records of D. acerinum from
other Acer spp. are probably all referable to other Drepanosiphum species,
e.g. aceris, caucasicum. Sutakova (1984) studied associations with microbial
organisms in relation to transmission of maple leaf perforation virus, and
Keller (1987) recorded a fungal infection.

Drepanosiphum aceris Koch Alatae yellowish or pale whitish-green with

dark thoracic lobes, siphunculi dark at least distally and often with rather
narrow cross-bands on ABD TERG 4 and 5, plus lateral spots in front of bases
of siphunculi and dark spots at wing-tips; BL 2.7-4.2 mm. On Acer campestre,
usually under leaves near ground, in Europe and east to the Caucasus. Sex-
uales in September-November.

Drepanosiphum braggii Gillette Alatae pale yellow to greenish with yellow-

brown thoracic lobes and dark-tipped siphunculi; BL 2.7-3.6 mm. On Acer
negundo, undersides of leaves, in western USA and Mexico (BMNH colln,
leg. R. Peña). Sexuales in September-November (Palmer, 1952). 2n = 30.

Drepanosiphum caucasicum Dzhibladze Alatae pale yellow with dark dorsal

abdominal markings on ABD TERG 4 and 5, or 5 only, very like D. acerinum
but with siphunculi whitish transparent, only the extreme apices being some-
times slightly dusky; BL 2.4-2.9 mm. On Acer trautvetteri in Georgia
(Dzhidbladze, 1962).

Drepanosiphum dixoni Hille Ris Lambers Alatae greenish-ivory with brown

head, dark brown to black thoracic lobes, pale brown to black siphunculi,
and two extensive blackish sometimes coalescing cross-bars on ABD TERG
4-5, plus dark sclerites in front of siphuncular bases; BL 2.5-3.3 mm.
Brachypterous alatae are common in midsummer (Dixon, 1972b). On Acer
campestre, under lower leaves in shade, sometimes on water-shoots growing
from lower parts of trunk. Recorded from England, Netherlands, former
Yugoslavia and northern Italy (van Harten and Coceano, 1981). Sexuales in
October (Hille Ris Lambers, 1971).

Drepanosiphum iranicum Hille Ris Lambers Colour of alata in life not

recorded, probably pale green, with variably pigmented head and thorax, and
with a conspicuous black spot in front of base of each siphunculus (even in
pale specimens); BL 3.1-4.1 mm. On Acer cappadocicum and A. velutinum
in Iran (Hille Ris Lambers, 1971). Oviparae in October (BMNH colln, leg.
S. Hodjat). Described as a subspecies of, and closely related to, D. platanoidis,
but the constancy of abdominal pigmentation and the different host relation-
ship indicate that this is a good species. 2n = 30*.
 Dysaphis 665

Drepanosiphum oregonensis Granovsky Alatae have orange-brown thoracic

lobes and pale green abdomen; BL 1.9-2.6 mm. Darker specimens may have
a blackish-brown mesosternum and dark spots in front of bases of the rather
thick, swollen, dark-tipped siphunculi. The black stripe on the fore femur
and black basal section of ANT III are distinctive. On various Acer spp.
in southern and Central Europe, Mediterranean, southwest Asia, India
(Chakrabarti, 1988), and introduced to western USA (California, Oregon).
Sexuales in October in California, but oviparae collected in-July in Germany
(BMNH colln, leg. W. Quednau). Michelena Saval and Gonzalez Funes
(1988b) recorded a parasitoid in Spain.

Drepanosiphum platanoidis (Schrank), Plate 7a Alatae have yellow-brown

head and thorax with darker brown markings and a pale green abdomen with
or without variably developed dark cross-bars (but if present these are never
restricted to ABD TERG 4-5); BL 3.2-4.3 mm. On undersides of leaves of
Acer pseudoplatanus; recorded from many other Acer spp., but these are
apparently only visited on a casual basis. It seems to be common on sycamores
wherever they are grown in Europe, Central Asia, North Africa, Australia,
New Zealand, USA and Canada (British Columbia). Sexuales in September-
November. There has been much work on this well-known aphid, including
studies of its population dynamics (e.g. Chambers et al., 1985; Wellings et al.,
1985), distribution on leaves (Dixon, 1976), seasonal variation (Dixon, 1972d,
1974, 1975), function of siphunculi (Dixon and Stewart, 1975), effects on
wood formation (Dixon, 1971a), flight behaviour (Dixon and Mercer, 1983),
sexual morph determination (Dixon, 1971b), effects of predation by ants
(Warrington and Whittaker, 1985) and chemical control (Parry et al., 1989).
2n = 30.

DYSAPHIS Börner Aphidinae: Macrosiphini

A palaearctic genus of about 110 species, many of which are heteroecious

with spring colonies distorting and discolouring the leaves of their primary
hosts in the Pyroidea (Amelanchier, Crataegus, Malus, Pyrus, Cotoneaster,
Sorbus), before migrating to secondary hosts in Umbelliferae and certain
other families. On the primary host they are generally medium-sized, rather
plump-bodied dark-legged aphids, greenish, bluish or pinkish-grey in colour
and covered in wax meal. Characteristic features are the short, often helmet-
shaped CAUDA and usual presence of spinal tubercles on head and posterior
abdominal tergites. Colonies are almost always attended by ants. In Dysaphis
s. str. on Malus, the fundatrix forms a small apical primary gall on the
young leaf, but her progeny soon leave this to form and develop in marginal
leaf-rolls. Migration of many heteroecious species occurs in the second
and/or third generation, so the spring colonies are short-lived and may
666 Dysaphis

comprise only the immediate descendants of the fundatrix; but in subgenus

Pomaphis colonies may persist for several generations on the primary host.
Shaposhnikov (1956, 1986) and Stroyan (1957a, as Sappaphis; 1963, 1985)
have published extensively on Dysaphis. Shaposhnikov and Moralev (1979)
reviewed and keyed the Crataegus-feeding species, and regional reviews are
available for the Indian subcontinent (Chakrabarti and Medda, 1993) and
Fennoscandia and Denmark (Heie, 1992). The common apple- and pear-
feeders are reviewed in B & E, 1984, pp. 263-268.

Dysaphis affinis (Mordvilko) Living without host alternation on Malus

spp. (orientalis, sieversii) in southwest and Central Asia (Armenia, Georgia,
Tadzhikistan, Turkey) and also recorded from Latvia and western Siberia
(probably introduced on seedlings). Injurious to apple tree nurseries, roll-
ing and blistering the leaves (Shaposhnikov, 1964a). The second genera-
tion includes many alatae as well as apterae. Sexuales occur in October
(Shaposhnikov, 1986). 2n = 12.

Dysaphis angelicae (Koch) Forming cherry-red curled-leaf galls on Cra-

taegus. Fundatrix is greenish-grey. All or nearly all the second generation are
alatae, and migrate to Angelica spp., forming colonies on lower leaf bases
(Stroyan, 1963). In Europe (England, France, Germany, Hungary, The
Netherlands, Sweden). (Note: spring migrant alatae of a closely-related species
that migrates to Laserpitium, D. laserpitii (q.v.), may key out to angelicae.)

Dysaphis anthrisci (Börner) A complex of sibling species and/or host races

rolls the margins of the leaves of apple in spring in Europe, the separate
taxa being only reliably distinguished by their life cycles and/or secondary
host relationships. Dysaphis anthrisci s. str. occurs throughout Europe and
migrates in the second generation to the leaf bases of Anthriscus sylvestris,
gynoparae and males returning to Malus in October. Spring populations of
D. devecta, D. chaerophylli and D. radicola (q.v.) in Europe are all similar
to those of anthrisci and only distinguishable with some difficulty. In the
former USSR, Shaposhnikov (1956, 1959, 1965, 1990) recognized other
closely-related apple-feeding taxa: D. anthrisci ssp. majkopica migrating from
apple to Anthriscus nemorosa; D. chaerophyllina, D. brachycyclica and
D. caucasica going to Chaerophyllum spp.; D. bunii to Bunium spp.; D.
physocaulis to Physocaulis nodosus; and D. flava and D. zini with unknown
secondary hosts. Shaposhnikov (1964a, 1965) provided keys to distinguish
some of the taxa in this very difficult species complex, which was discussed
further by Stroyan (1985). Members of the anthrisci group also occur in East
Asia, e.g. in Korea on Malus baccata (see Stroyan, 1985), and in China (Zhang
and Zhong, 1984a). 2n = 12.

Dysaphis apiifolia petroselini (Börner) On undersides of leaves of Cra-

taegus, forming cherry-red to crimson curled-leaf galls. Fundatrix is greenish-
grey, immature alatae greenish. The second generation are all alatae and
migrate to various Umbelliferae, e.g. Petroselinum, Conium, Apium (see
 Dysaphis 667

Stroyan, 1963). In northern and Central Europe. Dysaphis apiifolia s. str. is

the name conventionally applied to anholocyclic populations on Umbelliferae
that occur in many parts of the world (see B & E, 1984, p. 264). However,
red galls agreeing with apiifolia s. str. have been found on Crataegus azarolus
in Italy (Stroyan, 1985), indicating that this form may also have at least a
partial host alternation. Shaposhnikov (1987a) gave a key to the species of the
petroselini,group. 2n = 12.

Dysaphis (Pomaphis) ariae (Börner) In leaf-roll galls on Sorbus aria in

alpine Europe (Austria, France). Apterous fundatrigeniae were straw-white
(Börner, 1950). Stroyan (1963) considered that this might be a starvation form
of plantaginea, but it appears to be a distinct, shorter-haired species (see also
Shaposhnikov, 1963). It is possible that D. gallica on Linaria cymbalaria is
the secondary host form of D. ariae, although host transfers were so far unsuc-
cessful (Stroyan, 1985).

Dysaphis armeniaca Shaposhnikov Rolling and reddening the lateral edges

of the leaves of Malus pumila and M. orientalis in Armenia. Immatures are
pinkish-grey (cf. D. foeniculi malidauci). Second generation almost all alatae,
migrating to an unknown secondary host; of numerous attempted transfers
to various Umbelliferae only Ligusticum alatum elicited a settling response,
so the true secondary host may be a related species (Shaposhnikov, 1986).

Dysaphis (Pomaphis) aucupariae (Buckton) Spring generations live in con-

spicuous rolled- or twisted-leaf galls on Sorbus torminalis, which are initially
pale yellow-green, becoming tinged with red and later golden yellow. Despite
Buckton's name for this aphid, it occurs on Sorbus aucuparia only rarely.
Migration occurs in the third and subsequent generations to Plantago spp.
(Stroyan, 1957a). In Europe, east to Crimea and Caucasus; also presumably
anholocyclic populations are found on Plantago in Australia and New Zealand
(Sunde, 1984), and alatae have also been trapped in Washington, USA
(BMNH colln; leg. S.H. Halbert). 2n = 12.

Dysaphis brachycyclica Shaposhnikov Rolling leaves of Malus spp. in Russia

(northern Caucasus, Crimea, Armenia) in spring, but the galls are yellow
rather than red (cf. chaerophyllina, etc.). All second generation are alatae,
migrating in May to Chaerophyllum bulbosum. Return migration in July-
August. Shaposhnikov (1990) provided a detailed comparison with the closely
related D. chaerophyllina, A member of the D. anthrisci species complex.

Dysaphis brancoi (Börner) In deep purplish-red leaf-roll galls on Malus

pumila or M. orientalis in spring, migrating in second and third generations
to Valeriana spp., where it lives at or near ground level. Throughout Europe
(except Iberian peninsula), Kazakhstan, India (Kashmir; BMNH colln)
and Siberia (Pashchenko, 1988b). Populations in Britain tend to be longer-
haired and have been separated as a subspecies (D. brancoi rogersoni;
Stroyan, 1963).
668 Dysaphis

Dysaphis chaerophylli (Börner) Causing leaf-roll galls on Malus in spring.

Second generation almost entirely alate, migrating to Chaerophyllum spp.
(aromaticum, hirsutum, temulum, but perhaps not bulbosum; Stroyan, 1963).
In Western and Central Europe (Austria, the Czech and Slovak Republics,
England, Germany, Switzerland). A member of the D. anthrisci species group.

Dysaphis chaerophyllina Shaposhnikov Rolling and reddening the edges of

the leaves of Malus pumila and M. orientalis in Russia (northern Caucasus,
Armenia, Crimea), migrating in second and third generations (May-June) to
Chaerophyllum maculatum and, to some extent, Ch. bulbosum. Return
migration from Ch. maculatum in September-October. Shaposhnikov (1990)
provided a detailed comparison with the closely-related D. brachcyclica, and
Stekolshchikov and Lobanov (1990) computed discriminant functions for all
morphs of these two species. A member of the D. anthrisci species complex.

Dysaphis crataegi (Kaltenbach) Forming deep cherry-red curled-leaf galls on

Crataegus spp. in spring. Fundatrix greenish-grey, immature alatae pinkish.
Second generation nearly all alate, migrating to ground level parts of
Umbelliferae. A complex of closely related species or "subspecies" (host races)
occurs with different primary and/or secondary host plant relationships
(Stroyan, 1963, 1985; Shaposhnikov and Moralev, 1979). In Western Europe,
D. crataegi s. str. preferentially colonizes Daucus carota; ssp. kunzei (Börner)
goes to Pastinaca; and ssp. aethusae (Börner) to Aethusa cynapium and Torilis
japonica. In southwest Asia, ssp. pallida Shaphoshnikov and Moralev forms
typical red galls containing greenish-white immatures (cf. incognita) on
Crataegus songorica, but its secondary host has not been established, and ssp.
heraclei Shaposhnikov and Moralev migrates from Crataegus orientalis to
Heracleum pastinacifolium. In fact, all these taxa are part of a larger group
of Crataegus-feeding and host-alternating species that includes angelicae,
apiifolia, incognita, laserpitii, lauberti, ranunculi and virgata (subgenus
Crataegaria of Shaposhnikov, 1964a). These are treated separately here;
however, the classification as either subspecies of crataegi or as separate
species is somewhat arbitrary, not being based on any valid biological criteria.
Populations regarded as crataegi s. str. on Daucus occur in southwest Asia
and USA, where they are probably anholocyclic on the secondary host plant.
2n = 12 (on Daucus).

Dysaphis devecta (Walker) Rolling and reddening the edges of the leaves of
Malus in spring; the galls contain a mixture of normal bluish-grey wax-
powdered apterae and dark green to reddish alatiform apterae with different
degrees of pigmentation and sclerotization of head and thorax. Monoecious,
with an abbreviated life cycle of only three-four generations; oviparae and
alate males are produced within the galls (Hille Ris Lambers, 1945; Forrest,
1970). Forrest and Dixon (1975) studied gall formation, Llewellyn and
Hargreaves (1986) studied the energetics of gall-feeding, and Alston and
Briggs (1977) investigated the genetic basis of resistance to this aphid in apple.
Hitherto restricted to Europe, but Zhang et al. (1990a) record this species from
 Dysaphis 669

China. Despite its unusual biology, comparable morphs are very similar to
those of D. anthrisci and its relatives, and indicate a close relationship to that
group.

Dysaphis flava Shaposhnikov Rolling lateral margins of leaves of Malus

pumila in spring, and migrating in the second generation to an unknown
secondary host. Immatures in the leaf-rolls are pale yellow (Shaposhnikov,
1956). In European Russia and Kazakhstan. A member of the anthrisci group.

Dysaphis incognita Shaposhnikov and Moralev Forming red curled-leaf

galls on leaves of Crataegus curvisepala and Crataegus sp. in Crimea and
Armenia. Second generation all alate, migrating to an unknown host (transfer
experiments to many potential hosts were unsuccessful - see Shaposhnikov
and Moralev, 1979). Immatures in the Crataegus galls are yellowish or
brownish-pink (cf. D. crataegi ssp. pallida). See also Stroyan (1985).

Dysaphis laserpitii (Börner) Spring generaions on Crataegus sp(p).

undescribed, but presumably forming similar galls to other members of the
crataegi group. Secondary host is Laserpitium latifolium. Recorded from
Germany, France and Sweden. Gynoparae sometimes produce numerous
oviparae on Crataegus sp. in autumn (BMNH colln, leg. H.L.G. Stroyan).
Spring migrant alatae not available and therefore this species is not included
in the key under Crataegus. It is closely related to C. angelicae and C. lauberti
(Stroyan, 1963), and specimens could key to either of those species. (Alatae
collected in July on Laserpitium in Sweden have 0-10 secondary rhinaria on
ANT V and longest hairs on ABD TERG 2-3 are 22-36 µm long and rather
abruptly pointed.)

Dysaphis lauberti (Börner) Forming red blistered-leaf galls on Crataegus

leaves in spring, with second generation alatae usually migrating to form sum-
mer colonies on lower leaf-bases and root collar of Heracleum sphondylium,
although there are records from other Umbelliferae (Stroyan, 1963). Return
migration of gynoparae and males in September. In Europe, and southwest
and Central Asia (Turkey, Tadzhikistan; BMNH colln). (Note: spring migrant
alatae of the closely related species laserpitii (q.v.) may key to lauberti.)

Dysaphis malidauci Shaposhnikov Rolling and reddening the lateral margins

of leaves of Malus in spring, migrating in the second generation to Daucus
(Shaposhnikov, 1987b). Immatures in the leaf-rolls are green. In Kazakhstan.
Described as a subspecies of foeniculus Theobald, which occurs throughout
the world as anholocyclic populations on Umbelliferae (especially Daucus,
Foeniculum) and Polygonaceae (Rumex), but which is not known to have a
sexual phase. Dysaphis malidauci is here regarded as a separate species.
2n = 12.

Dysaphis mordvilkoi Shaposhnikov Rolling and reddening the lateral

margins of leaves of Malus (pumila, pallasiana) in spring in Siberia (Amur,
670 Dysaphis

Primorsk). Life cycle not studied, but suspected secondary host is Libanotis
seseloides (Shaposhnikov, 1986).

Dysaphis (Pomaphis) parasorbi (Börner) Forming leaf nests on Amelan-

chier ovalis in Austria (Börner, 1952-3). Life cycle unknown. Very similar to
D. sorbi, except for the very short antennal hairs (those on ANT III less than
0.2 x diameter of segment). See Stroyan (1963).

Dysaphis (Pomaphis) pavlovskyana Narzikulov Spring generations distort

and discolour the leaves of Sorbus spp. (persica, turkestanica) in Tadzhikistan
(Narzikulov, 1957). Secondary host is possibly Plantago sp. (Shaposhnikov,
1974; Stroyan, 1985). Similar, but probably not identical, aphids have been
collected on S. graeca in Turkey (G. Remaudière, cited by Stroyan, 1985)
and a subspecies, indica, has been described from S. cuspidata in India
(Chakrabarti and Medda, 1993). Khuda Bukhsh and Pal (1983a) recorded
2n = 12 for an aphid identified as this species and collected in India on Pyrus
?lanata.

Dysaphis physocaulis Shaposhnikov Rolling and reddening the leaves of

Malus pumila in Crimea, migrating in early May to Physocaulis nodosus, with
return migration in June after only one to two generations on the secondary
host (Shaposhnikov, 1990). A member of the D. anthrisci species complex.

Dysaphis (Pomaphis) plantaginea (Passerini) Forming yellowish crumpled-

leaf galls on Malus spp. and sometimes on Pyrus in warmer regions. Spring
migrant alatae appear in the third generation and migrate to form colonies
along veins on undersides of leaves of Plantago spp. In Europe, North Africa,
southwest and Central Asia, India, Pakistan, Nepal and North America. Also
recorded from Plantago only in Japan, Korea and Thailand. Recent work on
this well-studied aphid includes population dynamics (Graf et al., 1985),
interactions with hypersensitive apple plants (Lyth, 1985) and timing of
control measures (Hull and Starner, 1983). See also B & E, 1984, p. 266.
2n = 12.

Dysaphis (Pomaphis) pyraria Narzikulov Spring generations distort the

leaves of Pyrus korshinskyi in Tadzhikistan. Life cycle unknown; only the
apterous fundatrigenia is described (Narzikulov, 1961c; Stroyan, 1963;
Shaposhnikov, 1988). Closely related to D. pyri.

Dysaphis (Pomaphis) pyri (Boyer de Fonscolombe) Spring generations

distort and yellow the leaves of Pyrus communis; also recorded from
P. amygdaliformis, P. nivalis and P. ussuriensis. Apterae are wax-covered,
brownish-red to dark brown with black siphunculi. Migration to Galium spp.
occurs after two to three generations on pear, which in some years can result
in heavy attacks. In Europe, North Africa, southwest and Central Asia, nor-
thern India, Nepal and Pakistan. Santas (1987) recorded predators of colonies
on P. amygdaliformis in Greece. To judge from a paratype aptera in the
 Dysaphis 671

BMNH collection, D. multisetosa Basu, described from P. communis in West

Bengal, seems to be a synonym of pyri. See also B & E, 1984, p. 267 and
Stroyan (1957a).

Dysaphis radicola (Mordvilko) Spring generations roll and redden the

lateral margins of leaves of Malus (orientalis, pumila), in the same way as
others species of the anthrisci group. However, the small primary gall near
the apex of the leaf produced by the fundatrix comprises a longitudinal fold
near the mid-rib, rather than the transverse fold made by fundatrices of
anthrisci, etc. (Shaposhnikov, 1964a). Alatae are produced in the second
generation and migrate to form colonies on the roots of Rumex spp.
Anholocycly on Rumex roots also occurs commonly where this species has
never been found on Malus; e.g. in Britain (Stroyan, 1963) and Australia
(ACT; BMNH colln, leg. VFE). It occurs throughout Europe, east to Iran,
and alatae apparently of this species have been trapped in USA (BMNH colln).
Shaposhnikov (1964a) described a shorter-haired form with similar host rela-
tions in Russia (Crimea, Caucasus) as a subspecies, D. radicola meridialis and
this was given separate species status by Eastop and Hille Ris Lambers (1976).
Dysaphis radicola-group populations in Greece, Spain and Portugal are
similarly short-haired. 2n = 12.

Dysaphis ramani Das and Raychaudhuri Only apterous viviparae are

described, as dark brown, wax-covered aphids heavily infesting and curling
leaves of Crataegus sp. in April in Nepal (Das and Raychaudhuri, 1983).
Presence of apterae without alatae indicates that colonies persist on the
primary host beyond the second generation, which is unusual for a Crataegus-
feeding species of Dysaphis. Life cycle unknown.

Dysaphis ranunculi (Kaltenbach) Curled-leaf galls on Crataegus are pale

yellowish-green, often suffused with rosy-pink, but never with the sharp
demarcation line between red of gall and green of leaf lamina found in
crataegi. Fundatrix deep blue-grey, with a grape-like bloom, immature alatae
brownish to grey. Migrating in second generation to basal parts of Ranunculus
spp., returning to Crataegus in September (Stroyan, 1963). In Europe and
Central Asia.

Dysaphis (Pomaphis) reaumuri (Mordvilko) Colonies roll and twist both

leaf laminae and petioles of Pyrus spp. Apterae are light green covered with
wax in early spring, later dark green, with pale siphunculi. Migration occurs
to Galium spp., but colonies also persist on pear and even produce oviparae
in autumn (Kolesova, 1974). In southern Europe and Central Asia. 2n = 12.
See also B & E, 1984, p. 268.

Dysaphis sharmai Stroyan Appearance in life unknown; on Pyrus pashia in

Nepal, presumably distorting leaves. Also in India (Himachal Pradesh;
BMNH colln, leg. DHRL). Life cycle unknown, but presence of apterae
indicates that the species is either monoecious on Pyrus or migrates to a
672 Elatobium

secondary host in third or later generations (Stroyan, 1982). Shaposhnikov

(1988) discussed the taxonomie affinities of this aphid.

Dysaphis sibirica Shaposhnikov Galls on Malus in form of 'tuberculate

swellings of dark red colour' (Shaposhnikov, 1986, quoting other authors). All
second generation alate, migrating in mid-June, possibly to Carum carvi
and Sphallerocarpus gracilis (Umbelliferae) and/or Galeopsis bifida (see
Shaposhnikov, 1986). In eastern Siberia.

Dysaphis (Pomaphis) sorbi (Kaltenbach) Forming crumpled-leaf galls in

spring on Sorbus spp., especially aucuparia. Mid-rib and leaflets curl into tight
bunches, with only slight discoloration. Alatae are not produced in large
numbers until the fourth generation on Sorbus in June; they migrate to Cam-
panula spp. and perhaps to Jasione montana. Host alternation seems to be
facultative; colonies may persist on Sorbus into late summer and possibly these
produce sexuales without host alternation (Stroyan, 1957a; Shaposhnikov,
1963). Throughout Europe except the Iberian peninsula, and east to Turkey
and Caucasus. 2n = 12.

Dysaphis sorbiarum (Narzikulov) Apterae pale greenish-yellow, slightly

wax-dusted; reported to live on upper surfaces of leaves of Crataegus altaica
and C. hissarica in Tadzhikistan (Narzikulov, 1954). Apparently monoecious
on Crataegus, with alate males, alate females being rare (see also Stroyan,
1963).

Dysaphis (Pomaphis) tschildarensis Daniarova Spring generations distort

the leaves of Pyrus communis in Tadzhikistan and Kazakhstan. Host alter-
nation occurs to Ferula spp. and Archangelica decurrens (Shaposhnikov,
1988).

Dysaphis virgata Shaposhnikov and Moralev Forming pale green or red

curled-leaf galls on Crataegus orientalis in Armenia (Shaposhnikov and
Moralev, 1979). Immatures in galls are rose-pink with grey pubescence.
Second generation alate, migrating to an unknown secondary host. See under
crataegi, and Stroyan (1985).

ELATOBIUM Mordvilko Aphidinae: Macrosiphini

A genus with five biologically diverse species. Three are conifer feeders,
one lives on Salix and one on Trochodendron. The Japanese species were
revised by Miyazaki (1971), although his concept of the genus is broader
including related aphids (Ericolophium, Neoacyrthosiphon) that live on
Ericaceae.
 Elatobium 673

Elatobium abietinum (Walker), Plate 14a, b Apterae pale green, with two
darker green longitudinal stripes; BL 1.0-2.0 mm. Feeding on the needles of
Picea spp. and, much less commonly, Abies spp. In Europe, Tasmania, New
Zealand (Zondag, 1983a), Chile (Carrillo, 1977), Falkland Islands and western
North America (e.g. Koot, 1983). Monoecious holocyclic (with alate males)
in continental Europe (von Scheller, 1963), apparently anholocyclic elsewhere.
Spring colonies on Picea spp., especially P. sitchensis, cause discoloration
and loss of old needles, sometimes involving serious defoliation (Parry,
1969). Overwintering survival was studied by Powell and Parry (1976) and
flight activity by Carter and Cole (1977). Variation in susceptibility to
attack by E. abietinum has been studied between species of Picea (Nicholls,
1987) and between provenances (Day, 1984; Carter and Nichols, 1988) and
individual trees (Day, 1986) of P. sitchensis. Specialized parasitoids are
Lysaphidus schimitsheki (Stary, 1974) and Ephedrus koponeni (Halme, 1992).
2n = 18.

Elatobium hidaense (Takahashi) Apterae milky white, slightly pale

brownish at apices of antennae and legs; BL c. 1.5mm. On an unidentified
Salix sp. in Japan (Takahashi, 1961c) and also recorded from S. udensis in
Kamschatica (Pashchenko, 1988b). Described in Cavariella and possibly it
should be regarded as an aberrant member of that genus. Another Salix feeder
described in Elatobium, E. salicifoliae Zhang from Salix sp. in China, is said
to resemble E. hidaense but has a shorter, broader R IV+V (Zhang and Zhong,
1985a).

Elatobium laricis (Rupais) Apterae broadly oval, green, shining, without

wax, similar in colour to larch needles; BL 1.5-1.8 mm. On Larix sibirica in
eastern Siberia (Rupais, 1974). Other morphs and life cycle unknown.

Elatobium momii (Shinji) Apterae green: BL 0.9-1.6 mm. On needles of

Abies firma in Japan and also on Picea jezoensis, if the synonymy with
Neomyzaphis piceana Inouye is correct (Takahashi, 1960e). Shinji (1941)
also records Taxus cuspidata as a host. Sexuales and life cycle unknown.
(Specimens of an Elatobium sp. from Abies bornmuelleriana and A. nord-
manniana in Turkey (BMNH colln, leg. H. Çanakçioglu) have an antennal
PT of similar length to that of momii, but seem to be another, undescribed
species.)

Elatobium trochodendri Takahashi Apterae pale yellowish-green, with a

dark green thorax, a large dark green dorsal abdominal patch and siphunculi
that are black on distal two thirds; BL 1.3-2.2 mm. On Trochodendron
aralioides in Japan, feeding on upper sides of leaves along mid-ribs
(Takahashi, 1960e).
674 Eonaphis

EONAPHIS Essig Greenideinae: Cervaphidini

Four African species associated with Euphorbiaceae, differing from Schoute-

denia in the presence of spinal processes on the head, thorax and anterior
abdominal segments. Remaudière (1988) revised the genus.

Eonaphis crotonis Quednau Apterae broadly oval, somewhat flattened, dirty

brownish-yellow, shining, with a diffuse brown patch on dorsal abdomen, pale
finger-like dorsal processes and dark brown conical siphunculi; BL 1.1-
1.7mm. Alatae have forewing veins dark-bordered, the media being once-
branched. In large colonies on leaves and young branches of Croton
subgratissimus, especially along mid-ribs on lower leaf surfaces, attended by
ants (Quednau, 1962). In South Africa. Sexuales and life cycle unknown.

Eonaphis phyllanthi Remaudière Apterae mainly pale, with antennae dark

beyond the midpoint of ANT HI, and dark conical siphunculi; BL c.
1.1-1.2 mm. On Phyllanthus sp. in Madagascar (Remaudière, 1988). Other
morphs and life cycle unknown.

EPIPEMPHIGUS Hille Ris Lambers Pemphiginae: Pemphigini

A genus of four or five poplar gall-forming species, resembling Pachypappa

in that the fundatrices have no wax pore-plates and the spring migrant alatae
have secondary rhinaria with ciliated rims, but also with some characters of
Pemphigus, e.g. alatae have forewings with unbranched media and the first
instar exules have short empodial hairs. Chakrabarti and Banerjee (1993)
reviewed the Indian species.

Epipemphigus imaicus (Cholodkovsky) Galls on Populus ciliata are elon-

gate, caterpillar-like, reddish-green, 4-7 cm long, on upper surface of leaf
usually alongside the mid-rib, with an opening on underside (Fig. 131B;
A.K. Ghosh et al., 1981; Maity and Chakrabarti, 1981). The second generation
are alatae (BL 2.0-2.6 mm), emerging from galls in June-July and migrating
to form colonies on roots of Polygonum alatum (Chakrabarti and Banerjee,
1991, 1993). Sexuparae return to poplar in November-December (Chakrabarti
et al., 1985). A record of galls of E. imaicus on Ailanthus glandulosa
(Chakrabarti et al., 1971) must be in error and a record from the roots of Pinus
wallichiana (Habib and Ghani, 1970) was possibly a misidentified Prociphilus.
In Pakistan and northern India. Pemphigus chomoensis Chang and P. yun-
nanensis Chang, described from Populus sp. in China (Chang and Zhong,
 Eriosoma 675

1979c), are very similar and possibly both synonyms of E. imaicus. 2n = 18

(Khuda-Bukhsh, 1980).

Epipemphigus marginalis Chakrabarti and Banerjee Galls on Populus

ciliata in northern India are greenish-yellow to yellowish, 2.5-3.0 cm long,
produced on the marginal area of the leaf lamina. Second generation are all
alatae, BL 1.7-2.2 mm, migrating in May-June to found colonies on roots of
Impatiens falcifer (subject to experimental confirmation). Alate sexuparae
return to P. ciliata in September-October. Chakrabarti and Banerjee (1993)
described all morphs.

Epipemphigus niisimae (Matsumura) Galls on Populus spp. (koreana,

maximowiczii, suaveolens) are elongate, caterpillar-like or cockscomb-like,
yellow to crimson-red, along mid-rib of leaf on upper surface with opening
underneath (Aoki, 1975). Second generation are all alatae (BL 2.1-2.4 mm),
emerging from galls in June-July. Secondary host is not definitely known,
possibly Agrimonia pilosa and/or Bidens tripartita (see Aoki, 1975). Aoki and
Makino (1982) studied gall usurpation and lethal fighting between the heavily-
armoured 1st instar fundatrices. In Japan, Korea and Siberia. 2n = 20
(Blackman, 1986).

Epipemphigus sanpupopuli (Chang and Zhong) Galls on Populus cathayana

are elongate cockscomb-like, adjacent to mid-rib on upper surface of leaf,
opening underneath (Fig. 131 A; Zhang and Zhong, 1983b). Alatae, BL 2.0-
2.2mm, were collected in May (Chang and Zhong, 1979c). Secondary host
unknown.

ERIOSOMA Leach Pemphiginae: Eriosomatini

About 30 species, most of which have host alternation between galls on Ulmus
and secondary hosts in Pyroidea, Grossulariaceae or Compositae. Feeding by
the immature fundatrix on a growing elm shoot in spring induces a galling
reaction in one or more leaves distal to the feeding site, the fundatrix moving
into the leaf-roll after it is formed (Akimoto, 1981). Typically, alatae and
apterous exules of Eriosoma have rather conspicuous siphuncular pores with
partially chitinized rims and surrounding hairs. Alatae usually have a once-
branched media in the forewing, but specimens of E. yangi can have both
forewings with unbranched media.
The limits of the genus are difficult to define; the East Asian Colophina
and the North American Georgiaphis are closely related and often included
as subgenera (Smith, 1985; Eastop, 1987), but are treated separately in
this book. Eriosoma s. str. comprises four or five North American species
forming rosette leaf galls on Ulmus spp. of the americana group, with
676 Eriosoma

summer generations in woolly masses on Pyroidea, plus one species that is

monoecious and free-living on Ulmus. Among the palaearctic species, Eastop
(1987) distinguished two groups or subgenera; Schizoneura that typically form
open, curled-leaf galls on Ulmus and then migrate to roots of Grossulariaceae
and other plants, and Mimaphidus that typically form closed, bloated-leaf
galls on Ulmus before migrating to roots of Pyroidea, or in one case, Senecio.
These two groups are approximately equivalent to the ulmi group and
lanuginosum group discussed by Akimoto (1983), but there are some discre-
pancies. The life cycles of several species are still unknown. Areopraon
lepelleyi is a specialized parasitoid of the elm-feeding generations.
There are accounts or partial revisions of Eriosoma from Sweden
(Danielsson, 1979), India (A.K. Ghosh, 1984b), Siberia (Pashchenko, 1988b),
Japan (Akimoto, 1983) and North America (Smith, 1985). The species having
Ribes as secondary host were reviewed by Danielsson (1982). See also B & E,
1984, pp. 269-271.

Eriosoma americanum (Riley) Spring colonies on Ulmus americana roll the

edges of the leaves downwards. Emigrant alatae emerging from galls in June
are dark green to blue-black with some wax wool, and rather dark wing veins;
BL 2.0-2.5 mm. They migrate to form colonies of pale pink, woolly apterae
on roots of Amelanchier. Return migrants to elm in August to October pro-
duce dwarf sexuales that lay overwintering eggs on bark (Patch, 1915a;
Palmer, 1952). It occurs throughout the natural range of U. americana in
eastern North America, and in the west on planted elms.

Eriosoma (Schizoneura) anncharlotteae Danielsson Galls formed on Ulmus

(carpinifolia, procera) by inward curling of both edges of leaves, which
become blistered, yellow or light green, or sometimes bright red on younger
trees and often considerably distorted (Danielsson, 1979). Fundatrices dark
green to bluish-grey covered in wax meal. Alatae mainly produced in third
generation are dark bluish-grey, BL 1.8-2.3 mm, migrating in June-July to
roots of Ribes alpinum. Sexuparae fly back to elm bark from mid-September.
In Europe (the Czech and Slovak Republics, England, Germany, Poland,
Sweden) and also recorded from China (Zhang et al., 1985).

Eriosoma (Schizoneura) auratum Akimoto Forms a leaf-roll gall on Ulmus

japonica; one edge of the leaf is rolled tightly downward, and becomes
yellowish-green and stiff (Akimoto, 1983). Fundatrix is greyish-green. Almost
all second generation are alatae, brown, BL 1.5-1.9 mm, migrating in June
to found colonies on roots of Anthriscus sylvestris (Akimoto, 1985b). In
Japan and Korea. Gall invasion and competition with other Eriosoma on
Japanese elm were studied by Akimoto (1988a, 1989). Eriosoma eligulatum
Pashchenko (1988b), described from U. japonica and U. pumila in the eastern
Siberia, is possibly a synonym. 2n = 12.

Eriosoma crataegi (Oestlund) Apterae on bark of twigs and stems of

Crataegus spp. are bluish-black covered in white mealy wax and often have
 Eriosoma 677

two long wax filaments extending from the end of the abdomen; BL
2.0-3.0 mm. Widely distributed in North America. Crataegus is the usual host,
but aphids apparently of this species are also sometimes collected on bark of
Fagus, in colonies of Grylloprociphilus imbricator (Smith, 1985). Smith
distinguished several forms of apterous exules on Crataegus, differing in
dimensions of wax glands. Sexuparae fly to Ulmus (americana, fulva) and
produce sexuales on elm bark in September-October. Heriot (1938) described
differences in the sexuparae, males, oviparae and eggs between E. americana,
E. lanigerum and E. crataegi. Leaf rosette galls similar to those of E. lanigerum
are presumably formed on Ulmus in spring, but strangely, despite the
biological studies of Heriot (1938) and Cox (1939), the spring generations of
crataegi on elm have not been specifically identified; separation of crataegi
in the Ulmus key is based on data provided by Cox for spring migrant alatae
collected on Crataegus in May-June. 2n = 12.

Eriosoma (Mimaphidus) flavum Jancke (= E. gomboriense) Forming leaf-

roll galls on Ulmus spp. (carpinifolia, glabra, procera); the leaf tissue is hyper-
trophied and blistered between the veins (Fig. 134N, p. 898) and turns pinkish-
brown. Alatae, BL 1.4-1.7 mm, emerge in mid-July-August. Summer genera-
tions are on the roots of Pyrus communis and Cydonia oblonga, often deep
underground, the apterous exules being very small (BL 1.0-1.2 mm) and
yellowish (cf. pyricola), with filamentous wax. In Europe (England, The
Netherlands), Israel (exules on Cydonia), Iran (BMNH colln, leg. S.H. Hodjat)
and Georgia (as gomboriense; Dzhibladze, 1965a).

Eriosoma (Schizoneura) grossulariae (Schule) Forming open, green to

yellow curled-leaf galls on Ulmus spp. (Fig. 134L, p. 898), similar to those of
E. ulmi, with which this species was confused for many years (see Danielsson,
1982). Immature alatae in galls are light greyish-green (cf. ulmi), maturing in
June; dark green when adult, migrating to found colonies on roots of Ribes
sanguineum and R. uvacrispa. Throughout Europe and also introduced on
European elms into North America (B & E, 1984, p. 270). 2n = 10.

Eriosoma (Schizoneura) harunire Akimoto Leaf-roll galls on Ulmus

japonica are formed by rolling one margin of the leaf near its tip firmly
towards the underside and remain green and unswollen even when mature
(Akimoto, 1981). Fundatrix is brownish or greenish-grey and produces both
apterous and alate fundatrigeniae. Alatae mature in the galls in July, later
than E. auratum (Akimoto, 1989) and migrate to found colonies on roots of
Plantago asiatica (Akimoto, 1985b). Only known from Japan. Gall invasion
and competition with other Eriosoma were studied by Akimoto (1988a, 1989).
2n = 10.

Eriosoma (Schizoneura) japonicum Matsumura Leaf-roll galls on Ulmus

japonica are indistinguishable from those of E. harunire (see above).
Fundatrix reddish-brown. Alatae emerge in June and migrate to found
colonies on roots of Rosaceae (Agrimonia pilosa, Geum japonicum, Fragaria
678 Eriosoma

ananassa; Akimoto 1983, 1985b). In Japan, China (Zhang et al., 1990a) and
Taiwan (Tao, 1970; as E. ulmosedens). 2n = 10.

Eriosoma (Schizoneura) kashmiricum L.K. Ghosh, Verma and Raychaudhuri

Forming leaf-roll galls on Ulmus laevigata and U. wallichiana; the leaf margin
is folded downwards and twisted into a spiral (A.K. Ghosh et al., 1981).
Fundatrix is dark brown (Bhattacharya et al., 1980). Alatae are produced in
June and fly to an unknown secondary host. 2n = 12 (Pal and Khuda-Bukhsh,
1983).

Eriosoma (?Schizoneura) laciniatae Pashchenko Forming curled-leaf galls

on Ulmus laciniata in Siberia (Pashchenko, 1988b). Alatae are produced in
July, secondary host (if any) unknown. Appears closely related to grossulariae
or longicornutum, but lacks a ciliary fringe to the primary rhinarium on ANT
V. Some fundatrices collected from yellow curled leaves of U. laciniata in
China were possibly of this species (BMNH colln, leg. VFE, No. 18,182); these
specimens had 2n = 16*, different from any other known karyotypes of
Schizoneura.

Eriosoma lanigerum (Hausmann) Apterae on Pyroidea are purple, red or

brown, covered with thick white flocculent wax; BL 1.2-2.6 mm. On roots,
trunk or branches, often causing deformation and cancer-like swellings of
bark. A severe pest of apple (e.g. Weber and Brown, 1988). Anholocyclic
populations, sometimes with an apparently abortive sexual phase, occur on
apple throughout the world, overwintering on the roots (see B & E, 1984,
p. 270). In North America, aphids of the E. lanigerum complex induce leaf
rosette galls on Ulmus americana, but there has been doubt about the relation-
ship of these to apple-feeding populations. Smith (1985) successfully trans-
ferred alatae from such galls to apple. However, alatae indistinguishable from
those of E. lanigerum were obtained from reddish-pink leaf curls on U. alata,
and these would not transfer to apple but successfully colonized Crataegus
uniflora. Similar aphids occur in leaf-cluster galls on sterile elms in Australia
(Fisk et al., 1992).
Ecology and natural enemies of E. lanigerum on apple were most recently
studied by Bouchard et al. (1984; Quebec, Canada), Thakur et al. (1988; India)
and von Kogler (1989; Germany). Aphelinus mali is a widespread specialized
parasitoid (but Aphelinus gossypii, which parasitizes a wide range of hosts
including Aphis and Rhopalosiphum spp., have often been misidentified as
A. malí). A study of predation by earwigs was made by Mueller et al. (1988).
J. Davidson (1913) made a detailed study of morphology and anatomy. See
Molinari (1986) for a general review. 2n = 12.

Eriosoma (Mimaphidus) lanuginosum (Hartig), Plate 5e Forms clusters of

large, closed, bloated-leaf galls near ends of branches of various Ulmus spp.
(Fig. 134M, p. 898). The gall is produced by extreme hypertrophy of the leaf
parenchyma on one side of the mid-rib near its base; the hypertrophied tissue
is light green at first and covered with fine white hairs, becoming brown as
 Eriosoma 679

the gall matures (Marchal, 1933). Fundatrices are blackish, wax-powdered,

apterous fundatrigeniae are dark green with black legs. Alatae produced in
the second and third generations on elm are dark green to black, wax-
powdered; BL 2.1-3.1 mm. They emerge from the galls in late June-July and
migrate to found colonies on fibrous rootlets of Pyrus communis or Cydonia.
Apterae on pear roots are variable in colour, pale yellow to reddish, BL
2.0-2.7 mm. Return migration to elm bark occurs in September. Occurs
throughout Europe, the Mediterranean area and southwest Asia. Often con-
fused with E. pyricola, a smaller species with more caudal hairs which has a
much wider geographical distribution (de Fluiter, 1933; B & E, 1984, p. 271).
2n = 10.

Eriosoma (Schizoneura) lishanense Akimoto Forms leaf-roll galls on Ulmus

uyematsui in mountains of Taiwan. Fundatrix has 5-segmented antennae.
Emigrant alatae are similar to those of E. auratum, with secondary rhinaria
distributed III 23-26, IV 5, V 0, VI 0, and BL 1.4-1.5 mm (Akimoto, 1983).
Life cycle and secondary host unknown.

Eriosoma (Schizoneura) longicornutum Akimoto Forms open leaf-roll galls

on Ulmus japonica and U. laciniata in Japan; the leaf edge on one side is rolled
downwards and becomes inflated and yellowish-green. Fundatrix yellowish-
green with dark brown legs and antennae. Alatae produced in June-July
migrate to an unknown host. Closely related to E. grossulariae, but with a
difference in the wax glands of first instar exules (Akimoto, 1983). 2n = 10.

Eriosoma mimicum Hottes and Frison On Ulmus spp. (americana, fulva) in

eastern North America, probably forming leaf-curl galls. Alatae collected in
June, migrating to an unknown host (Hottes and Frison, 1931; Smith, 1985).
Georgiaphis maxsoni is probably a synonym.

Eriosoma (Schizoneura) moriokense Akimoto Forms leaf-curl galls on

Ulmus japonica; often whole leaves are strongly twisted, becoming reddish or
brownish and dusted with wax powder when the gall is mature (Akimoto,
1983). Fundatrix is reddish-brown and produces both apterous and alate
fundatrigeniae. Alatae (BL 1.6-2.2 mm) leave galls in late June and found
colonies on roots of Sedum spp. (Akimoto, 1985b). Sexuparae return to trunk
of elm in September. In Japan, China (BMNH colln, leg. VFE) and probably
Korea; according to Akimoto, Eriosoma sp. B of Paik (1972) is probably
E. moriokense. Akimoto (1983) records evidence of aggressive behaviour
towards predators by second and third instar larvae in galls. 2n = 10.

Eriosoma (Mimaphidus) patchiae (Börner and Blunck) Galls consist of

curled, twisted and blistered leaves on shoots of Ulmus spp. (glabra, montana,
procera), stunting and twisting young growth, often on suckers arising from
the trunk (Marchal, 1933). Galls green or yellowish, fundatrices yellowish-
green, apterous fundatrigeniae whitish-green, according to Marchal (1933);
but fundatrix is dark bluish-grey according to Danielsson (1979). Alatae
680 Eriosoma

produced in the second and subsequent generations are green or brownish,

with narrow brown dorsal abdominal cross-bands; BL 2.0-2.5 mm. Migration
occurs in June-July to roots of Senecio, or Cineraria, with return to elm in
September-October. However, the host alternation seems to be facultative,
because alatae emerging from galls later in July-August are often sexuparae.
Possibly a distinct, monoecious species is involved, for which the name
ulmosedens Marchal, 1919 is available, but according to Marchal (1933) some
of the alatae from galls can give birth to both sexuales and apterous exules.
In northwest and Central Europe. Parker (1984) studied the association of
E. patchiae with Anthocoris gallarum-ulmi, and Dessart and Gardenfors
(1985) studied parasitism and hyperparasitism. Eriosoma dilaniginosum Zhang
(1980b), described from U. pumila in China, is very similar and possibly a
synonym. 2n = 10*.

Eriosoma (Schizoneura) phaenax (Mordvilko) Leaf galls on Ulmus (densa,

montana, procera, wallichiana) appear to be similar to those of patchiae; the
illustration of a gall-cluster on U. campestris (= procera) by Mordvilko (1924,
Pl. IV), ascribed to patchiae and copied by Shaposhnikov (1964a) and Heie
(1980), applies to phaenax. The simple marginal leaf-roll described for
E. phaenax by A.K. Ghosh et al. (1981) was perhaps a young gall. Apterous
fundatrigeniae in galls are light brown (A.K. Ghosh, 1984b). Emigrant alatae
emerging from galls in June-July are brownish with narrow darker brown
dorsal abdominal cross-bands; BL 2.1-2.5 mm. Apterous exules have not been
described, but sexuparae have been collected in October on Cotoneaster in
Pakistan (BMNH colln, leg. R.H. Ghani) and Kashmir, India (BMNH colln,
leg. Rishi), and this is probably the secondary host; sexuparae collected on
Androchne cordifolia (Euphorbiaceae) in Pakistan (A.K. Ghosh, 1984b) were
probably vagrants. Mordvilko (1935), however, noted that E. phaenax was
monoecious, so possibly host alternation is facultative, as in E. patchiae. In
Transcaucasus, Turkmenia, Tadzhikistan, Afghanistan, Pakistan and India.
This species was at one time erroneously synonymized with patchiae (see
Marchal, 1933; Narzikulov, 1965a).

Eriosoma (Mimaphidus) pyricola Baker and Davidson Galls on Ulmus spp.

(americana, procera) are clusters of bloated leaf tissue like those of
E. lanuginosum (see Alfieri, 1920; as inopinatum). Fundatrix and apterous
fundatrigeniae grey, emigrant alatae produced in second and third generations
are brown with pale yellow abdomen (cf. lanuginosum); BL 1.3-2.4 mm.
Migration occurs in June-July to fibrous rootlets of Pyrus communis or
Cydonia. Apterous exules are yellowish-pink when immature, pink to red
when adult, rather sparsely clothed with filamentous wax (Baker and David-
son, 1916); BL 1.3-2.0 mm. Sexuparae are shiny dark green to brown, and
return to elm bark in September-October. In southern Europe (Italy,
the former Yugoslavia), Turkey, and introduced into southeastern Australia,
Tasmania, New Zealand, USA (western and some northeastern states),
Canada (British Columbia), South America (Argentina, Chile) and (probably)
South Africa. Swenson (1971) studied the influence of the secondary host on
 Eriosoma 681

sexupara production. This aphid is often confused with E. lanuginosum in

the literature. Eriosoma alabastrum Pashchenko (1988b), described from
U. japonica in Siberia, is very similar and possibly a synonym.

Eriosoma rileyi Thomas On bark of trunk and branches of small elms

(Ulmus americana, fulva) in North America, forming small colonies covered
with wax wool very like those of E. lanigerum on apple and causing similar
deformation of woody tissue. Monoecious holocyclic on elm, without forming
leaf-galls. Apterae under the wax are pinkish, yellow or brown; BL 1.8-
2.3 mm. Alatae produced in June-August have ANT III 0.8-1.0 x head width
across (and including) eyes, with secondary rhinaria rather widely spaced.
Sexuparae are apterous, produced in September-October, depositing dwarf
sexuales directly onto bark (Patch, 1913b; Palmer, 1952).

Eriosoma (Schizoneura) sorbiradicis Danielsson Apterae yellowish-white,

with sparse wax; BL 0.7-1.4 mm. They live singly or in small colonies on
rootlets of Sorbus aucuparia (Danielsson, 1979). Recorded from Scotland
(Stroyan, 1991), Sweden (Danielsson, 1979) and Oregon, USA (BMNH colln;
leg. R.L. Furniss). Other morphs and life cycle unknown.

Eriosoma (Schizoneura) ulmi (Linnaeus) Galls on Ulmus spp. are formed by

downward-curling, twisting and blistering of one lateral edge of a leaf, the
deformed part becoming yellowish- or whitish-green. Fundatrix and apterous
fundatrigeniae are wax-covered, dark green, immature alatae are brownish
and wax-covered (cf. grossulariae). Adult alatae are dark green to bluish-grey,
migrating in June-July to found colonies on roots of Ribes rubrum and
R. nigrum (Danielsson, 1982). Return migration of sexuparae to elm bark is
in September-November. In Europe (including Iceland and Faroes), southwest
and Central Asia, to Mongolia and China according to Heie (1980). Eriosoma
mediocornutum Pashchenko (1988b), described from Ulmus japonica and
U. pumila in Siberia, is very similar and possibly a synonym. Parker (1984)
studied predation by Anthocoris gallarum-ulmi, and Dessart and Gardenfors
(1985) studied parasitism and hyperparasitism. (B & E, 1984, p. 271.) 2n = 10.

Eriosoma (Schizoneura) ulmipumilae Ivanoskaya In curled-leaf galls on

Ulmus pumila in the Altai region of Siberia (Ivanoskaya, 1976). Life cycle
unknown. Closely related to E. japonicum and E. yangi.

Eriosoma (Schizoneura) yangi Takahashi Forms yellowish-green rolled-leaf

galls on Ulmus parvifolia in Japan, China and Korea (Akimoto, 1983). Alatae
from galls in May-June have BL 1.4-1.7 mm, and migrate to found colonies
on roots of Salix sp. and Fragaria ananassa (Akimoto, 1985b). A closely-
related form regarded as a subspecies, E. yangi parasiticum, occurs on Ulmus
japonica in Japan and is an obligatory 'cuckoo' parasite of other Eriosoma
species, invading and usurping their galls; Akimoto (1988a, b) made a detailed
study of this phenomenon and compared morphology and behaviour of
fundatrices of the two forms.
682 Essigella

ESSIGELLA Del Guercio Lachninae: Cinarini

About 14 nearctic species, all living on Pinus needles except one species
(wilsoni) specific to Pseudotsuga. They are long-bodied aphids, as befitting
their needle-feeding habit, and are the nearctic equivalent of Eulachnus, but
smaller and with 5-segmented antennae. Apterae are spindle-shaped, with
tapered abdomen, but the 'spindle' is truncated at the head end, which is
relatively broad. They are often overlooked, as they feed near the bases of
the needles and move quickly when disturbed; their biology is consequently
little known. Sorensen (1988, 1992, 1994) revised Essigella, synonymizing
many of the species described by Hottes (1957c, 1958f).

Essigella alyeska Sorensen Apterae spindle-shaped, grey-green with yellow-

orange head and mainly pale appendages; BL 1.4-1.7 mm. Feeding singly on
needles of Pinus banksiana in Canada (Ontario, Quebec) and also collected
on Picea glauca in Alaska (Sorensen, 1988). ABD TERG 2-4 have 8
spinopleural and 2 marginal hairs, and ABD TERG 8 has 6-8 hairs. Alatae
have 0-2 secondary rhinaria on ANT III. Other morphs unknown.

Essigella braggi Hottes Apterae spindle-shaped, grey-green, or grey to light

brown; BL 2.2-2.5 mm. On needles of Pinus contorta (var. murrayana) in
Sierra Nevada and Cascades of California, USA (Hottes, 1957c; Sorensen,
1994). Closely related to E. knowltoni, and regarded as part of the knowltoni
species complex by Sorensen (1992).

Essigella californica (Essig), Plate 16c Apterae spindle-shaped, with grey-

green thorax and lime green abdomen, with or without brown dorsal spots;
BL 1.5-2.0 mm. Legs variably pigmented, often mainly pale, tibiae some-
times darker. Alatae have 2-4 secondary rhinaria on ANT III. Feeding singly
on needles of Pinus spp. of subsections Cembrae, Strobi, Ponderosae,
Sabinianae, Oocarpae and several introduced palaearctic pines, but rarely
found on pines of subsections Contortae or Cembroides, and records from
such plants are quite likely to be misidentifications of other species. It also
sometimes occurs on Pseudotsuga (menziesii, macrocarpa). In western North
America from southern British Columbia and Alberta, south to Mexico and
introduced into Europe (France and Spain; Turpeau and Remaudière, 1990;
Seco Fernandez and Mier Durante, 1992). Anholocyclic in California; life
cycle at more northerly latitudes unstudied. 2n = 8.

Essigella critchfieldi Sorensen Apterae spindle-shaped, black, or rarely

green with black dorsal spots; BL 1.6-1.9 mm. Feeding on needles of Pinus
contorta s. str. (with one record from var. latifolia) in coastal northwestern
Canada and USA as far south as northern California (Sorensen, 1994).
 Essigella 683

Essigella eastopi Sorensen Apterae spindle-shaped, varying in colour from

wholly yellow to dark brown with yellow on front of head and a yellow spinal
stripe on thorax and abdomen; BL 1.6-2.0 mm. On needles of Pinus coulteri
in coastal ranges of California, south of San Francisco Bay, to Mexico
(Sorensen, 1994).

Essigella essigi Hottes Apterae spindle-shaped, black or dark green through-

out, or with yellow-green head and green thorax and abdomen, often with
dark dorsal spots; BL 1.6-1.8 mm. On needles of Pinus spp. of subsection
Oocarpae (attenuata, radiata), and rarely on subsection Ponderosae. In
southwest Oregon and California.

Essigella fusca Gillette and Palmer Apterae spindle-shaped, head and

thorax dusky yellow-brown, abdomen green with brown spots; BL 2.0-
2.6mm. Feeding singly on needles of Pinus spp. of subsection Ponderosae,
especially P. ponderosa; also recorded from P. coulteri and P. leiophylla, and
collected once from Callitris drummondii (BMNH colln, leg. H.G. Walker).
In southern British Columbia, Canada and south through Rocky Mountains
to Mexico. Apterous males and oviparae occur in September-October
(Palmer, 1952). Populations in California and southwest Oregon are regarded
as a distinct subspecies (voegtlini) by Sorensen (1994).

Essigella hillerislambersi Sorensen Apterae spindle-shaped, straw-yellow,

often with dark spots, with head yellow to orange-brown; BL 2.0-2.7 mm. On
needles of Pinus jeffreyi in southwest Oregon and California, USA (Sorensen,
1994).

Essigella hoerneri Gillette and Palmer Apterae spindle-shaped, head and

thorax grey-green, abdomen lime green, legs yellow-brown; BL 1.4-2.0 mm.
On needles of Pinus spp. of subsection Cembroides (pinyon pines) in south-
western USA; a common species, and the only Essigella to preferentially
colonize pinyon pines (Sorensen, 1994). Oviparae occur in September-
November. Closely related to, and difficult to distinguish from, E. californica,
with which it has frequently been confused in the literature.

Essigella kathleeni Sorensen Apterae narrowly spindle-shaped, wholly pale

yellow; BL 1.3-2.0 mm. On needles of Pinus lambertiana in southwest Oregon
and California, USA (Sorensen, 1988). Oviparae collected in September.
Other morphs unknown.

Essigella kirki Sorenson Apterae narrowly spindle-shaped, grey-green or

occasionally pale yellow; BL 1.7-2.1mm. On needles of Pinus flexilis and
P. strobifomis in the Rocky Mountains from Montana to Arizona and New
Mexico, and in the southern Sierra Nevada and White Mountains of Califor-
nia, USA (Sorensen, 1988). Oviparae collected in September, other morphs
unknown.
684 Eucallipterus

Essigella knowltoni Hottes Apterae spindle-shaped, usually dark brown to

black with front of head yellowish, sometimes grey-green; BL 1.8-2.1 mm. On
needles of Pinus contorta (var. latifoliae and var. murrayanae) in western
Canada (British Columbia, Alberta) and in interior parts of Washington and
Oregon, south through the Rocky Mountains to Utah and southern Colorado
(Hottes, 1957c; Sorensen, 1992). Essigella braggi on P. contorta contorta in
California is closely related and regarded as part of the knowltoni species com-
plex by Sorensen (1992).

Essigella pini (Wilson) Apterae spindle-shaped, mainly green to greenish-

yellow with yellow-orange to reddish-orange head and with rows of brown
dorsal spots on abdomen (Wilson, 1919a); BL 1.5-2.0 mm. On needles of
Pinus spp. of subsection Australes (strobus, taeda, virginiana) in eastern USA,
with one record from Canada (southern Quebec). Alatae have about 3 secon-
dary rhinaria on ANT III. Other morphs are unknown. Records from other
hosts and from western North America (e.g. Palmer, 1952) are based on mis-
identifications. Patti and Fox (1981) studied the seasonal occurrence of
E. pini on P. taeda in South Carolina, where viviparae are present during the
winter months.

Essigella wilsoni Hottes Apterae are spindle-shaped, wholly lime green; BL

1.1-1.8 mm. On needles of Pseudotsuga spp. (menziesii, macrocarpa) in
western Canada (British Columbia, Alberta) and in western USA south to
Mexico. Not recorded from Pinus. Other morphs unknown.

EUCALLIPTERUS Schouteden Drepanosiphinae: Phyllaphidini

Active, delicate insects on Tiliaceae, with all viviparae alate. The anal plate
is deeply cleft. Parasitoids specializing on Eucallipterus and related genera
such as Chromaphis, Myzocallis, Tinocallis and Tuberculatus are: Trioxys
pcllidus (emerging from pale mummies), Praon flavinode (forming 'tents'
under the dead aphids) and Aphelinus subflavescens (emerging from black
mummies).

Eucallipterus tiliae (Linnaeus), Plate 9f Alatae are pale yellow with black
markings, including lateral stripes on head and prothorax and two rows of
black dorsal abdominal spots; BL 1.8-3.0 mm. On undersides of leaves of
Tilia spp. in Europe, southwest and Central Asia, North Africa, and intro-
duced to North America and New Zealand. Monoecious holocyclic; apterous
oviparae and alate males occur in late August to October. Numerous aspects
of the biology of E. tiliae have been studied including population dynamics
(Dixon, 1971c; Barlow and Dixon, 1980; Heimbach, 1986); aggregation behav-
iour (Kidd, 1976); flight behaviour (Kidd, 1977); sexual morph production
 Euceraphis 685

(Dixon, 1972a); effect on tree growth (Dixon, 1971b; Llewellyn, 1972); suscep-
tibility of different Tilia species (Carter and Nichols, 1986); interactions with
predators in UK (Wratten, 1973; Glen and Barlow, 1980); and parasitoids in
USA (Hajek, 1986). Biocontrol has been used in California, USA (Zuparko,
1983). 2n = 10 (female), 2n = 8 (male); the female has 2 pairs of X
chromosomes and sex determination is X 1 X 1 X 2 X 2 /X 1 X 2 0 (R.L. Blackman,
unpublished data).
[Besides the well-known E. tiliae, there may be at least two other species
in this genus: (i) Specimens with a slightly longer ultimate rostral segment from
Tilia tomentosa in Greece, Turkey and Iran may represent an undescribed
species; (ii) Shinji (1933) described Therioaphis tilicola from Tilia japonica
and T. miqueliana in Japan. T. tilicola has not since been recognized (Higuchi,
1972); it seems more likely to belong in Eucallipterus, rather than in the East
Asian Tilia-feeding genus Tiliaphis.]

EUCERAPHIS Walker Drepanosiphinae: Phyllaphidini

A genus of six or more species living on Betula and Alnus. They are rather
large aphids with a relatively short antennal terminal process, knobbed
CAUDA and rounded anal plate. The alder-feeding aphids are somewhat
distinct from the birch-feeders. All adults apart from oviparae are winged,
active insects which secrete bluish-white wax, often in the form of tufts on
their legs. The birch-feeding species often have black dorsal abdominal mark-
ings, but these vary greatly according to season and are often absent in mid-
summer. They seem very host-specific, although adults fly a lot and may be
found on many plants besides their true hosts (but see Pashchenko, 1984). The
oviparae are large, apterous, yellowish to dark brown with dark dorsal
abdominal markings and with the posterior abdomen prolonged into a conical
'ovipositor'. Keys are available for Europe and North America (Blackman,
1977, 1988) and Siberia (Pashchenko, 1988b).

Euceraphis betulae Koch, Plate 10a This name is currently applied to a com-
plex of morphologically similar species with different host associations, distin-
guishable by karyotype differences. Adults of the nominate species feeding on
B. pendula (= B. verrucosa) have a pale green to pale yellow abdomen with
bluish-white wax and a very variable degree of black pigmentation; BL
3.0-4.2 mm. The head and thorax are normally black, both dorsally and ven-
trally, the legs and antennae variably pigmented, sometimes quite dark and
the dorsal abdomen may be unmarked or have transverse black bands (in
spring), or black patches on ABD TERG 3 and 4 only (in autumn). Oviparae
and alate males occur in September-November. Euceraphis betulae occurs
throughout Europe and on introduced B. pendula in North America, Australia
and New Zealand. A morphological comparison with E. punctipennis in
686 Euceraphis

Europe is given by Blackman (1977). Hajek and Dahlsten (1988) studied

ecology of introduced populations in California, and Neuvonen and Lindgren
(1987) studied effects of simulated acid rain in Scandinavia. Other members
of the complex occur in North America on B. occidentalis and B. papyrifera,
in arctic and subarctic regions on B. glandulosa/nana, and on several Betula
spp. in East Asia, e.g. B. ermanii and B. platyphylla var. japonica. Pash-
chenko (1984, 1988b) distinguished four species in Siberia; E. caerulescens
Pashchenko, E. betulijaponicae (Matsumura), E. ontakensis Sorin, and
'E. punctipennis', but the differences in pigmentation of legs and antennae
may be only locally applicable. Taking the world fauna as a whole, the
members of the betulae complex cannot be reliably distinguished except by
their karyotypes. 2n = 8, 9, 10 or 11 (for European and North American
species; Blackman, 1988).

Euceraphis gillettei Davidson Adult viviparae are active, pale green alate
aphids with pale brown thorax, dorsal abdomen never with dark markings and
only a small amount of wax; BL 2.5-3.5 mm. On Alnus spp. (rhombifolia,
rubra, rugosa, tenuifolia) in North America. Oviparae and alate males occur
in September-November. The name may be being applied to more than one
species, as indicated by the variation in karyotype. 2n = 15, 16, 18 or 19
(Blackman, 1988).

Euceraphis lineata Baker (= E. deducta Baker) Adults are pale yellow-

green, with distinctive brown and black dorsal markings; BL 3.3-4.0 mm. The
head and prothorax have a pale to dark brown longitudinal band, the
mesothoracic dorsal lobes are brown and in the most pigmented specimens the
scutellum may be black, but the thorax never has any pigment ventrally. Sex-
uparae in late summer-autumn usually have a large black patch on ABD
TERG 3 and a smaller one on ABD TERG 4. Femora are pale except for black
'knees', tibiae have a dark dorsal longitudinal stripe and black apices and tarsi
are black. Immatures have banded antennae and dark tarsi contrasting with
their pale tibiae. This species is specific to B. populifola in northeastern USA
and eastern Canada. Sexuales occur in September-October. 2n = 16.

Euceraphis mucida (Fitch) Adult viviparae are pale green to yellow-green

with an orange-brown thorax and dark tibiae coated with blue-grey wax; BL
3.4-3.8 mm. Dorsal abdomen usually unmarked in spring, but alate sexuparae
in late summer-autumn may have transverse bars on ABD TERG 4 and 5.
On B. lenta in western USA. Oviparae and alate males occur in September-
October. 2n = 20, 21 or 22, the differences in this case having no taxonomie
significance as they are attributable to variation in the number of accessory
(B) chromosomes (Blackman, 1988).

Euceraphis ontakensis Sorin Adult alate viviparae are pale green with dark
legs, often with black bars on ABD TERG 4 and 5 and secreting bluish-white
wax; BL 3.0-4.0 mm. On Betula ermanii in Japan; also recorded from Alnus
fruticosa in Japan (Sorin, 1970), and from several other Betula and Alnus spp.
 Eulachnus 687

in Siberia. The occurrences on Alnus may only be casual. Sexuales unknown.

2n = 22.

Euceraphis punctipennis (Zetterstedt) Adult viviparae are pale green with a

dark brown head and thorax, often somewhat larger than E. betulae (BL
3.0-4.8 mm) and with dorsal black patches when present confined to ABD
TERG 4 and 5, even in spring generations (Blackman, 1977). On B. pubescens
in Europe (including Iceland and the Faroes). Pashchenko's (1984) records
from Siberia probably apply to another member of the E. betulae group.
Oviparae and alate males occur in October-November. 2n = 7 or 8, according
to the number of accessory (B) chromosomes (Blackman, 1976).

EULACHNUS del Guercio Lachninae: Cinarini

About ten species of long-bodied aphids living on needles of Pinus, the

palaearctic equivalent of the nearctic genus Essigella. They are cryptic when
feeding, but very active when disturbed. The best-known species show prefer-
ences for certain Pinus spp., but none are strictly monophagous. Accounts are
available for Italy (Binazzi, 1983b, 1984, 1989), UK (Carter and Maslen, 1982),
Central Europe (Pintera, 1968; Szelegiewicz, 1978), former Czechoslovakia
(Pasek, 1954), Hungary (Szelegiewicz, 1978), India (A.K. Ghosh, 1982b),
Siberia (Pashchenko, 1988b) and Japan (Inouye, 1970). The genus needs fur-
ther revision, including biosystematic studies, as many of the characters that
have been used in species discrimination are undoubtedly subject to environ-
mental influences. Lampel and Burgener (1987) studied pylogenetic relation-
ships with other Cinarini based on enzyme electrophoretic data. Diacretus
leucopterus and Praon bicolor are specialized parasitoids.

Eulachnus agilis (Kaltenbach) Apterae are spindle-shaped, bright green with

numerous reddish-brown spots, without wax; BL 1.6-2.3 mm. The hind legs
often have mottled pigmentation. On many Pinus spp., but particularly com-
mon on P. sylvestris, usually feeding on old needles. It occurs throughout
Europe, east to Turkey, and on planted P. sylvestris in North America, where
feeding damage was studied by Bliss et al. (1973). Monoecious holocyclic;
oviparae and alate males are produced in October-November, and the
oviparae lay eggs singly, mainly on leaf scars on the branches (Bliss and
Kearby, 1971). Kearby and Bliss (1969) studied population trends and control,
Kidd et al, (1985) studied its association with Schizolachnus pineti.
Eulachnus agilis is a member of a closely related group of species discussed
and keyed by Binazzi (1983b, 1989). Eulachnus altica Börner, described from
P. mugo at high altitudes in Central and southeastern Europe (Heinze, 1962;
Pintera, 1968), will key to E. agilis in this book, and may or may not be
specifically distinct; the minor differences from agilis, involving hair lengths
688 Eulachnus

and pigmentation, could easily be due to geographical variation, or environ-

mentally induced. Gabrid (1989) records aphids identified as E. alticola from
numerous native and introduced Pinus spp. in Kirghizia. Eulachnus cretacea
Mamontova, from P. cretacea in Ukraine (Mamontova, 1968), is also very
close to agilis. Eulachnus intermedius Binazzi, described from P. mugo in a
montane locality in central Italy, may key to agilis, but is perhaps a short-
haired form of E. rileyi (q.v.). 2n = 8.

Eulachnus brevipilosus Börner Apterae spindle-shaped, bright green with-

out distinct brown dorsal spots and without wax; BL 1.4-2.2 mm. On needles
of Pinus spp. (mugo, nigra, sylvestris) throughout Europe and introduced
to New Zealand (Zondag, 1983b) and northwestern USA (Washington;
Hottes and Essig, 1955) and British Columbia, Canada (BMNH colln, leg.
C.K. Chan). Sexual morphs are unknown; viviparae have been collected in
winter months in England, indicating anholocycly. Ponsen (1981) studied the
digestive system. 2n = 30.

Eulachnus nigricola (Pašek) Apterae spindle-shaped, pale green, with

brownish legs and antennae and slight wax dusting on head, sides of thorax
and dorsal abdomen; BL 2.0-2.3 mm. On needles of Pinus nigra (incl. var.
pallasiana, var. salzmanii) in Europe (France, Spain, the Czech and Slovak
Republics, Hungary, Bulgaria), east to Turkey (BMNH colln). Life cycle not
studied, but oviparae are known, so presumably holocyclic (Pintera, 1968;
Szelegiewicz, 1978).

Eulachnus pumilae Inouye (= cembrae Börner) Apterae spindle-shaped,

bright green to yellowish-green; BL 2.3-3.0 mm. Dusted with wax powder
according to Pasek's (1954) description of cembrae, but wax is not mentioned
in the description of pumilae (Inouye, 1939). On needles of Pinus spp. of
subsection Cembrae (cembra, parviflora, pumila, koraiensis). In montane
Europe (Austria, Switzerland, Poland, the Czech and Slovak Republics),
northern India, Siberia, Korea and Japan. Oviparae and males occur in early
September in Europe (BMNH colln).

Eulachnus rileyi (Williams) (= bluncki Börner), Plate 16d Apterae spindle-

shaped, varying in colour from dark olive green to orange-brown or grey, with
a dusting of bluish-grey wax; BL 1.8-3.0 mm. At least the hind pair of legs
are usually dark. On needles of a taxonomically wide range of Pinus, including
many North American species, but on Old World pines of subsection
Sylvestres it occurs much more commonly on P. nigra and P. mugo than on
P. sylvestris. Heavily attacked needles turn yellow and are lost prematurely
(Felt and Bromley, 1936). In Europe, Mediterranean area, southwest Asia,
and introduced into Africa south of the equator (Mills, 1990) and North,
South and Central America. Monoecious holocyclic in Europe and North
America, with oviparae and alate males appearing in September-November,
but apparently anholocyclic in warmer regions (Marchant, 1981). Katerere
(1983, 1984) studied its population ecology in Zimbabwe and reported on the
 Eumyzus 689

incidence of a fungal infection. Tremblay and Micieli de Biase (1970) recorded

parasitism in southern Italy. 2n = 8.
Although most populations of E. rileyi are readily distinguished from the
related species E. agilis, forms with apparently intermediate or combined
characters of the two species occur in southern and Central Europe. Binazzi
(1989) described E. intermedius from P. mugo var. pumilio in the mountains
of central Italy, with the appearance in life of E. rileyi but hairs more like
those of agilis. Eulachnus tauricus Bozhko, described from Pinus nigra var.
pallasiana in Crimea and also recorded from Caucasus and Ukraine (Bozhko,
1961), also differs from typical E. rileyi only in the characters of the hairs on
the antennae and tibiae, and was given subspecies status by Hille Ris Lambers
(1966a). Populations on P. halepensis in Italy (Binazzi, 1983a) and on P. pinea
in Israel (Halperin, 1986) have subsequently been identified as E. rileyi
tauricus. However, without further biological information all these forms are
perhaps best regarded as geographic or environmentally-determined variants
of E. rileyi.

Eulachnus thunbergii (Wilson) Apterae are spindle-shaped, dirty green to

yellowish-green or yellowish-brown (A.K. Ghosh, 1982b); BL 2.3-3.2 mm. On
needles of various native Pinus spp. in East and Southeast Asia (India, China,
Japan, Korea, Siberia, Taiwan, Java, Philippines) and also in Australia
(New South Wales), where it was collected on P. taeda (Eastop, 1966).
Oviparae and alate males occur in Japan in October-November (Inouye, 1970)
and oviparae are also recorded from India (Raychaudhuri et al., 1983). Mao
(1986) studied population trends in Japan, and A.K. Ghosh (1982b) noted
predators and parasitoids in India. 2n = 8 (Khuda-Bukhsh and Kar, 1990).

Eulachnus tuberculostemmata (Theobald) Apterae are narrowly spindle-

shaped, pale green to greenish-yellow, with small brown dorsal spots; BL
1.4-2.7 mm. On needles of Pinus spp., especially halepensis (incl. var. brutia);
also canariensis, eldarica, pinaster, pinea and nigra (incl. var. maritima). In
southern Europe, Mediterranean area and southwest Asia. Biology is little
known and no sexual morphs are recorded. Eulachnus mediterraneus Binazzi
(1983a) differs only in characters that are strongly size-correlated, and without
biological information is best regarded as large tuberculostemmata. Aphids
described as E. tuberculostemmata ssp. garganicus Binazzi (1983b) seem to be
small specimens of the nominate species, probably due to the influence of
environment or host plant. 2n = 8.

EUMYZUS Shinji Aphidinae: Macrosiphini

About ten biologically diverse East Asian species, of which three described
from India roll the leaves of woody Rosaceae in spring. They are probably
690 Euthoracaphis

all heteroecious and holocyclic, but with unknown secondary hosts. Eumyzus
have a spinulose, Myzus-like head, but with divergent antennal tubercles, and
rather long stiff dorsal body hairs with tuberculate bases. Chakrabarti and
Bhattacharya (1985) revised the genus.

Eumyzus eastopi Maity and Chakrabarti Colour in life unrecorded, pro-

bably dark; BL of aptera 1.7-1.9 mm. On Pyrus vestita in June in Uttar
Pradesh, India, rolling the edges of the leaves (Maity et al., 1982; Chakrabarti
and Bhattacharya, 1985). Other morphs and life cycle unknown. 2n = 10
(Khuda-Bukhsh and Pal, 1986b).

Eumyzus pruni Chakrabarti and Bhattacharya Colour in life unrecorded,

probably dark, with dark siphunculi; BL of aptera 1.7-1.9 mm. On Prunus
cornuta in Uttar Pradesh, India, living in tight reddish marginal leaf-rolls in
spring (Medda et al., 1986). Heteroecious holocyclic, migrating to an unknown
secondary host; oviparae and alate males are described by Medda and
Chakrabarti (1989), collected on P. cornuta in November.

Eumyzus prunicolus Medda and Chakrabarti Colour in life unrecorded,

probably pale, with pale siphunculi; BL of aptera 1.8-2.1mm. On Prunus
padus in Uttar Pradesh, India, rolling the leaves in spring. According to
Medda and Chakrabarti (1986b), the fundatrices induce two types of gall;
either the edge of the leaf is curled ventrally and becomes somewhat shrunken,
shortened and rugose, or a caterpillar-like pouch gall is produced on the upper
side of the leaf lamina, with a slit-shaped aperture to the underside. Alatae
are produced in June and presumably migrate to an unknown secondary host.
Sexual morphs are unknown.

EUTHORACAPHIS Takahashi Hormaphidinae: Nipponaphidini

Small, heavily sclerotized aphids related to Nipponaphis, with rather numerous

fine hairs on dorsal and marginal areas of prosoma, and with sutures dividing
the dorsal area of the prosoma into a large quadrate central area, plus an
anterior and two dorsolateral areas. Three species; two on Lauraceae, and one
reportedly on Senecio (A.K. Ghosh and Raychaudhuri, 1973a; Noordam,
1991).

Euthoracaphis heterotricha Ghosh and Raychaudhuri Apterae are aleyrodi-

form, dark brown to bluish black with dorsum covered by a thin floury
secretion; BL 1.0-1.8 mm (Noordam, 1991). On lower sides of leaves of
Cinnamomum spp. (zeylandicum, iners) in Java. Originally described from an
unknown host in West Bengal (A.K. Ghosh and Raychaudhuri, 1973a). Alatae
in May. Sexual morphs and life cycle unknown.
 Eutrichosiphum 691

Euthoracaphis umbellulariae (Essig) (= cinnamoniae Shinji) Apterae are

aleyrodiform, sedentary, almost circular, pill-box-shaped, black, covered with
white filamentous wax (Essig, 1932); BL c. 0.8 mm. Alatae are jet black. On
undersides of leaves of Cinnamomum spp. (japonicum, camphoratum) in
Japan (Takahashi, 1959b), and presumably introduced from East Asia into
California, USA, where it is very common on Umbellularia californica and
also recorded from Sassafras variifolium (= albidum) and Cinnamomum spp.
(Essig, 1932; Hille Ris Lambers, 1966d). Sexuales and life cycle are unknown;
presumably any holocycle would involve Distylium as primary host. Popula-
tions in California are anholocyclic. Hille Ris Lambers (1966d) queried
Takahashi's synonymy of cinnamoniae and umbellulariae, but specimens in
the BMNH collection do not show any consistent differences between
Japanese and Californian populations.

EUTRICHOSIPHUM Essig and Kuwana

Greenideinae: Greenideini

A genus of nearly 40 species, mostly on Fagaceae; the exceptions are described

from Betulaceae, Elaeocarpaceae, Lauraceae, Apocynaceae, Sonneratiaceae,
Euphorbiaceae, Sterculiaceae and unidentified plants. Eutrichosiphum have an
elongate oval or pear-shaped body, a long, tapering and distinctly subdivided
last rostral segment, hind tibiae without stridulatory ridges, siphunculi without
any reticulation in apterae and a CAUDA without a median process. Separate
genera or subgenera have been erected for such aphids, grouping species
according to the number of antennal segments, extent of dorsal spinulosity and
the nature of the siphuncular or dorsal body hairs. None of these characters
is entirely satisfactory for subgeneric classification (A.K. Ghosh, 1987), so here
we adopt a conservative approach, regarding Brevitrichosiphon, Holo-
trichosiphon, Paratrichosiphum, Neoparatrichosiphum and Neotrichosiphum
as all part of Eutrichosiphum and not recognizing any subgenera. Raychaudhuri
(1956) revised the species then known, Takahashi (1962b) reviewed the Japanese
species, and Raychaudhuri and Chatterjee (1980) and L.K. Ghosh (1986)
provided accounts of the species of northeast India and Himachal Pradesh
respectively. A full account of the Indian species (B. Agarwala, pers. comm.)
and an account of the species of Java (D. Noordam) are in preparation.

Eutrichosiphum alnicola (Basu) Apterae brown (Das and Raychaudhuri,

1983); BL 1.8-2.7 mm, on tender shoots of Alnus nepalensis in northwest
India (A.N. Basu, 1968) and Nepal. Saha and Chakrabarti (1988a) described
the alate male, collected in October in western Himalaya. The alate oviparae
collected in association with greenish apterae on Quercus and ascribed to
alnicola by Raychaudhuri et al. (1980a) are unlikely to be this species. It is
not clear how alnicola differs from alnifoliae Tao (see below).
692 Eutrichosiphum

Eutrichosiphum alnifoliae (Tao) Apterae mainly blackish-brown; BL c. 1.7-

1.8mm. Alatae are light yellowish-brown. On tender shoots of Alnus
cremastogyne in Szechuan, China (Tao, 1958). Sexuales and life cycle
unknown. Eu. alnicola (Basu) could be a synonym. [Das and Raychaudhuri
(1983) described apterae of another species on Alnus nepalensis in Nepal under
the name Eu. alnifoliae. Their aphids were greenish, BL 3.2-3.5 mm, with a
longer antennal PT and longer R IV+V than alnicola Basu; their description
agrees remarkably well with that of Eu. betulae from Betula sp. in Sikkim,
India.]

Eutrichosiphum arunachali Basu, Ghosh and Raychaudhuri Apterae dark

brown; BL 1.3-1.6 mm according to original description, but 2.8 mm for one
specimen from the type series measured by Raychaudhuri and Chatterjee
(1980). Described from an unidentified Quercus sp. in Arunachal Pradesh,
India (R.C. Basu et al., 1972) and also recorded from Sikkim (Mondal et al.,
1979). The alata is described by Chatterjee et al. (1981); it is longer-haired than
that of the otherwise similar Eu. pasaniae.

Eutrichosiphum assamense Ghosh, Basu and Raychaudhuri Appearance in

life unrecorded, probably brown with pale antennae, legs and siphunculi; BL
of aptera 1.7-2.3mm (A.K. Ghosh et al., 1969a). On Quercus fenestrata in
Assam, India, and on Quercus sp., Nepal (BMNH colln, leg. K.C. Sharma).
Keyed in this book on the basis of the original description, plus two apterae
agreeing with that description from Nepal (but see Raychaudhuri and
Chatterjee, 1980). Other morphs and biology unknown. [This species is con-
fused with Eu. tattakanum ssp. assamense Ghosh and Raychaudhuri, 1964 in
the check list of Agarwala and Ghosh (1984), and most of the references they
give refer to the latter species, which has been synonymized with Eu. kha-
syanum (Raychaudhuri and Chatterjee, 1980).]

Eutrichosiphum betulae Mondal, Chatterjee and Raychaudhuri Appearance

in life not recorded, probably pale; BL of aptera 2.9-3.3 mm. On an uniden-
tified Betula sp. in Sikkim, India (Mondal et al., 1979). Other morphs and
biology unknown. To judge from the descriptions, Eu. alnifoliae Das and
Raychaudhuri, 1983 (nec. Tao, 1958) could be a synonym.

Eutrichosiphum davidi Raychaudhuri Apterae probably dark brown; BL

1.7 (holotype)-3.1 mm. On Quercus serrata in southern India (Raychaudhuri,
1956); also recorded from Quercus sp. in Sikkim (Mondal et al., 1979) and
alate viviparae have been trapped in Nepal (BMNH colln, leg. K.C. Sharma).
Sexuales and life cycle unknown.

Eutrichosiphum dubium van der Goot Apterae pear-shaped, dark brown;

BL 1.9-2.7 mm. On young growth of Quercus spp. (dealbata, glauca),
Castanopsis sp. and Lithocarpus spp. (glabra, konishii); also collected on
Litsea sebifera (Raychaudhuri et al., 1977). In India (West Bengal, Manipur),
Taiwan, Japan (BMNH colln, leg. RLB) and Hong Kong (BMNH colln, leg.
 Eutrichosiphum 693

D.S. Hill). Zhang and Zhong (1985f) distinguished a population from Yun-
nan, China (which included apterae with 5-segmented antennae) as spp.
yulongshanense. Devi and Singh (1987) studied ant attendance. 2n = 20.

Eutrichosiphum flavum Takahashi Apterae orange-yellow, with rather

short, pale siphunculi, pale legs, and ANT PT/BASE about 1; BL 1.6-
1.9mm. Some of the paratypes in the BMNH colln have 5-segmented anten-
nae. On young leaves of ?Lithocarpus sp. in Thailand (Takahashi, 1941);
also reported from Quercus dealbata and Quercus sp. in India (Manipur,
Meghalaya). Singh et al. (1980) described an alate vivipara purported to belong
to this species. Sexuales and life cycle unknown.

Eutrichosiphum garhwalense Maity and Chakrabarti Appearance in life not

recorded, probably rather dark; BL of aptera 1.8-2.0 mm. On Quercus incana
in Uttar Pradesh, India (Maity and Chakrabarti, 1980). Alate viviparae,
oviparae and alate males, all collected in June-July, were described by Saha
and Chakrabarti (1988a). Apterae of the type series include specimens with
both 5-segmented and 6-segmented antennae. This species is very similar to
Eu. simlaensis (described as having 5-segmented antennae) and apart from
having a somewhat shorter last rostral segment, both could be synonyms of
Eu. khasyanum (described with 6-segmented antennae).

Eutrichosiphum heterotrichum (Raychaudhuri) Apterae pear-shaped, dark

brown; BL 2.8-3.2mm. Described from Quercus sp. in Java (Raychaudhuri,
1956). Other morphs unknown. This species is very similar to, and perhaps
only a geographical variant of, Eu. dubium.

Eutrichosiphum jugeshwari Singh Apterae elongate pear-shaped, appear-

ance in life not recorded but probably rather pale, with black-tipped siphun-
culi; BL 2.3-2.6mm. On Quercus sp. in Manipur, India (Singh et al., 1979).
Other morphs unknown.

Eutrichosiphum khasyanum (Ghosh and Raychaudhuri) Apterae pear-

shaped, yellowish-brown to dark brown, with pale antennae, legs and siphun-
culi (Ghosh and Raychaudhuri, 1964); BL 1.8-2.1mm. [Raychaudhuri et al.
(1980a) reported cream-coloured apterae.] Originally collected from under-
sides of leaves of an unidentified Quercus sp. in Meghalaya, India; subsequent
records are from Qu. dealbata and Qu. griffithii in northern India and from
Qu. semicarpifolia in Nepal (Miyazaki, 1977). Raychaudhuri and Chatterjee
(1980) provide measurements of an alate vivipara identified as this species.
Specimens from Khasi Hills, northeast India on an unidentified oak have a
longer PT (PT/BASE ratio 1.54-1.93, compared with 1.09-1.30 in khasyanum)
and may be a distinct species (B. Agarwala, pers. comm.). Life cycle and sex-
uales unknown.

Eutrichosiphum litseae Raychaudhuri, Raha and Raychaudhuri Apterae

pear-shaped, colour in life unrecorded, probably mainly pale; BL 1.5-1.8 mm.
694 Eutrichosiphum

Described from Litsea sebifera in Manipur, India. Other morphs and biology
unknown. In every respect except host plant this species seems to resemble
Eu. flavum Takahashi.

Eutrichosiphum makii Raychaudhuri and Chatterjee Appearance in life not

recorded, probably dark brown; BL 1.5-1.7 mm. Described from Glochidion
sp. in Meghalaya, India (Raychaudhuri and Chatterjee, 1974) and subse-
quently found on Litsea sp. A.K. Ghosh (1976) described the alate vivipara.
Sexual morphs and biology unknown.

Eutrichosiphum manipurense Singh, Raychaudhuri and Raychaudhuri

Appearance in life not recorded, probably rather pale except for a brownish
central abdominal patch and dark-tipped siphunculi (T.K. Singh et al., 1979);
BL 1.7-2.1mm. On an unidentified Quercus sp. in Manipur, India. Other
morphs and biology unknown.

Eutrichosiphum nungsireiae Singh, Raychaudhuri and Raychaudhuri

Apterae pear-shaped, dark brown with paler head, antennae and legs and stout
blackish siphunculi; BL c. 1.3mm. On Quercus sp. in Manipur, India. Other
morphs and biology unknown. [Eu. mukerjee Raychaudhuri, M.R. Ghosh,
Banerjee and Raychaudhuri, of which the host is unknown, is a very similar
aphid, but with a shorter antennal processus terminalis, shorter last rostral
segment and shorter body hairs (Raychaudhuri et al., 1973).]

Eutrichosiphum pasaniae (Okajima), Plate 12d Apterae variably pigmented,

pale brown to dark greenish or blackish-brown with black siphunculi; BL
1.4-1.8 mm. Alatae have an extensive black dorsal abdominal patch. On
young shoots and undersides of leaves of Castanopsis spp. and Elaeocarpus
japonicus; it is likely that records from Quercus are erroneous. In East and
Southeast Asia. Alate males were recorded in Taiwan in October (Takahashi,
1923). Szelegiewicz (1968) distinguished populations on Castanopsis sp. in
Vietnam, with a pale body and longer last rostral segment than pasaniae from
Japan and Taiwan, as Eu. pseudopasaniae. However, there is little or no cor-
relation between pigmentation and length of last rostral segment in popula-
tions in northeast India, where both light- and dark-bodied apterae may occur
on the same plant (Raychaudhuri and Chatterjee, 1974). Eutrichosiphum
mengluense Zhang, 1980a, described from China, also seems to be inter-
mediate between pasaniae and pseudopasaniae. Eutrichosiphum izas Zhang,
only known from alate viviparae in China, is also in this group. Further work
is needed to establish whether or not a single, variable species is involved.

Eutrichosiphum pyri Chakrabarti, Ghosh and Raychaudhuri Apterae pear-

shaped, colour in life not recorded; BL 1.7-2.1 mm. Described from Pyrus
communis (Chakrabarti et al., 1972b), but presumably the true host is
Quercus; there are subsequent records from Qu. semicarpifolia and Quercus
sp. (Chakrabarti et al., 1988). In northern India (Meghalaya, Uttar Pradesh).
Other morphs and biology unknown.
 Eutrichosiphum 695

Eutrichosiphum quercifoliae Raychaudhuri, Ghosh, Banerjee and Ghosh

Apterae are elongate pear-shaped, pale (Raychaudhuri et al., 1973); BL
2.2-2.6 mm. Alatae have a solid brown patch on ABD TERG 2-4. Note that
the alate morph included in the original description was an ovipara, not a
vivipara (R.C. Basu and Raychaudhuri, 1980). Collected on Quercus sp. in
West Bengal in December. Other morphs and biology unknown. Eutricho-
siphum manoji Chatterjee, Mondal and Raychaudhuri, 1981, described from
Quercus sp. in Sikkim, seems to agree in nearly all respects with Eu. quer-
cifoliae, and is likely to be a synonym.
Eutrichosiphum querciphaga Chakrabarti and Maity Aptera pale except for
a dark brown patch of irregular shape on the dorsal abdomen; BL 1.8-2.1 mm.
On an unidentified Quercus sp. in Uttar Pradesh, India (Chakrabarti and
Maity, 1980). Other morphs and biology unknown.

Eutrichosiphum rameshi (Raychaudhuri, Chatterjee and Raychaudhuri)

Apterae pear-shaped, appearance in life unrecorded, probably pale with dark
brown siphunculi; BL c. 1.9mm. On Quercus dealbata in Meghalaya, India
(Raychaudhuri et al., 1977). Other morphs and biology unknown.

Eutrichosiphum raychaudhurii (Ghosh) Apterae pear-shaped, colour in life

unrecorded; BL 2.0-2.3 mm. Described from an unidentified Quercus sp. in
West Bengal, India (A.K. Ghosh, 1969). Specimens identified as this species
have subsequently been recorded several times from Alnus sp(p)., including
alate as well as apterous viviparae (Raychaudhuri and Chatterjee, 1974) and
an alate ovipara (collected in May; Singh et al., 1980). It is unlikely that these
Alnus-feeding aphids are the same species. According to the original descrip-
tion, the siphunculi of apterae of raychaudhuri have long and short hairs inter-
mingled over the entire surface, a characteristic feature of the subgenus
Holotrichosiphon. Biology unknown.

Eutrichosiphum russellae (Ghosh, Ghosh and Raychaudhuri) Apterae elon-

gate oval, dark green to blackish; BL 2.5-2.6 mm. Alate viviparae have
siphunculi about 0.33 x BL. Originally collected on young twigs of Quercus
sp. in West Bengal, India, attended by ants (A.K. Ghosh et al., 1971e); subse-
quently recorded from Q. dealbata in Meghalaya (Raychaudhuri et al., 1977).
Sexuales and life cycle unknown. Very similar to Eu. rameshi. Zhang and
Zhong (1985f) described apterous specimens from Quercus sp. in China as a
subspecies, russellae ssp. lijiangense Zhang.
Eutrichosiphum sankari Raychaudhuri, Ghosh, Banerjee and Ghosh Apterae
pear-shaped, probably rather dark; BL 1.3-1.9 mm. Described from an
unidentified host in West Bengal, India (Raychaudhuri et al., 1973); subse-
quently recorded on Litsea cubeba in Meghalaya by A.K. Ghosh (1976), who
described the alate vivipara. Sexuales and biology unknown.
Eutrichosiphum shiicola Takahashi Aptera elongate, yellowish, with siph-
unculi distally black; BL 2.3-2.5 mm. On Castanopsis cuspidata (principal
696 Eutrichosiphum

host) and Quercus dentata in Japan. Green and yellow alate males and alate
oviparae occur on Castanopsis in May-June in Osaka Prefecture, but not until
autumn in Tokyo (Takahashi, 1962b). Eutrichosiphum sclerophyllum Zhang
(1980a), described from Castanopsis sclerophylla in China, looks like a
synonym.

Eutrichosiphum sikkimense (Raychaudhuri, Ghosh, Banerjee and Ghosh)

Aptera elongate oval, colour in life not recorded, probably pale; BL
c. 2.1 mm. Described from a single aptera plus immatures collected on
Duabanga sonneratioides in Sikkim, India (Raychaudhuri et al., 1973). Pos-
sibly this is not the normal host plant.
Eutrichosiphum simlaensis L.K. Ghosh Aptera greenish with dark siphun-
culi; BL 1.7-2.0 mm. On undersides of leaves and apical shoots of an uniden-
tified Quercus sp. in Himachal Pradesh, India (L.K. Ghosh, 1986). From the
description this species is very similar to Eu. garhwalense.

Eutrichosiphum sinense Raychaudhuri Aptera elongate oval, yellow, with

siphunculi black distally; BL 1.5-1.8 mm. Alate viviparae have a brown dorsal
abdominal patch and black siphunculi a little more than 0.5 x BL (Takahashi,
1962b). Described from an unknown host in China (Raychaudhuri, 1956), and
subsequently recorded from Castanopsis cuspidata in Japan (Takahashi,
1962b; Moritsu, 1983), and from C. javanica in Indonesia (BMNH colln, leg.
D. Noordam). Apparently not found on Quercus, although Eu. narafoliae
(Shinji, 1922b), described from Qu. serrata in Japan and not since recognized,
seems closely related. Sexuales and life cycle unknown.
Eutrichosiphum subinoyi Raychaudhuri, Ghosh, Banerjee and Ghosh
Aptera pear-shaped, colour in life not recorded; BL c. 1.4mm. Alate vivipara
elongate, BL c. 2.0 mm, with siphunculi c. 1.0 mm. Described from Pterosper-
mum sp. in Meghalaya, India (Raychaudhuri et al., 1973). The description
of the aptera of this species agrees remarkably with that for Eu. takahashii,
collected in the same locality on Glochidion (see below); according to
Raychaudhuri and Chatterjee (1980), however, Eu. subinoyi has fewer dorsal
abdominal hairs (10-14 per segment) than related species.
Eutrichosiphum takahashii Basu, Ghosh and Raychaudhuri Aptera pear-
shaped, probably dark in life, with blackish siphunculi; BL c. 1.6mm. On
Glochidion sp. in Meghalaya, India (R.C. Basu et al., 1975) and also recorded
from Sikkim (Mondal et al., 1979). It seems very similar to Eu. subinoyi.
Other morphs and biology unknown.

Eutrichosiphum taoi Ghosh, Basu and Raychaudhuri Aptera elongate-oval,

colour in life not recorded; BL 2.2-2.3 mm. Described from an unidentified
plant in Meghalaya, India; subsequently apterae identified as this species were
collected on Quercus sp. in Uttar Pradesh (Chakrabarti et al., 1972b) and on
Q. serrata in Manipur (Agarwala et al., 1980). Other morphs and biology
unknown.
 Fimbriaphis 697

Eutrichosiphum tapatii Mondal, Chatterjee and Raychaudhuri Aptera

pear-shaped, pale green with dark markings on dorsal abdomen leaving a
pale central area, and dark siphunculi; BL 1.7-2.0 mm. Described from
an unidentified plant in Sikkim, India (Mondal et al., 1979); subsequently
apterae and immatures identified as this species were found on Quercus sp.
in Uttar Pradesh (Raychaudhuri et al., 1980a). Other morphs and biology
unknown.

Eutrichosiphum tattakanum (Takahashi) Aptera pear-shaped, darkish but

with a pale spinal area on thorax and anterior abdomen and pale siphunculi
with dusky apices (colour in life apparently unrecorded); BL 1.8-2.2 mm.
Alate viviparae have broad dark partially coalescent bars on ABD TERG 3-5.
On undersides of leaves of Quercus spp., Castanopsis formosana and also
Castanea formosana (Takahashi, 1931a). In Japan, Taiwan, northern India
and Nepal. Alate males and alate oviparae are produced in June-July in Hon-
shu (Takahashi, 1962b). Eutrichosiphum neoalnicola Raychaudhuri, Ghosh
and Das (1980a), described from Quercus sp. in India, is very similar and
possibly a synonym. Specimens from Q. dealbata and Castanopsis sp. in the
Khasi Hills, northeast India, have a long narrow R IV+V (0.38-0.43 mm,
and 3.17-3.58 x HT II, compared with 0.24-0.30mm and 1.92-2.50 x
HT II for tattakanum), and may be a distinct species (B. Agarwala, pers.
comm.).

FIMBRIAPHIS Richards Aphidinae: Macrosiphini

A genus of about ten mostly North American Myzus-like species, closely

related to the European genus Ericaphis and associated with Rosaceae,
Ericaceae and Liliaceae (see B & E, 1984, pp. 222-224). One species lives on
woody Rosaceae.

Fimbriaphis gentneri (Mason) Apterae yellow-green to green, broadly

spindle-shaped; BL 1.3-2.4 mm. Alatae have a broken dark dorsal patch. On
leaves and young shoots of various woody Rosaceae in western North America
(Amelanchier, Crataegus, Photinia, Sorbus and also on the non-native Pyrus
communis and Mespilus germanica). Monoecious holocyclic, with oviparae
and alate males appearing in September-October (BMNH colln, leg. C.K.
Chan). All morphs have been described, but the biology is little known. There
is considerable karyotype variation both within and between samples; 2n
(female) = 18, 19, 20, 21 and 23, and 2n (male) = 17 and 19, have all
been found in samples from British Columbia, Canada (R.L. Blackman and
C.K. Chan, unpublished data). The biological and taxonomie significance of
this variation are not yet understood.
698 Foaiella

FOAIELLA Börner Phylloxeridae

A genus for one phylloxerid which has apterae with fine hairs and alatae with
a round distal rhinarium, found on oak roots in southern Europe.

Foaiella danesii Grassi and Foá Apterae greenish-yellow, pear-shaped, with

broad head and thorax and tapered abdomen; BL 0.5-1.5 mm. Alatae have
a pale yellow head and a yellow costal margin and pterostigma in the forewing.
On roots of Quercus robur, which become tuberose. Holocyclic, producing
apterous and alate sexuparae in August-October. The sexuales lay overwinter-
ing eggs on the roots and possibly also in bark crevices on the trunk (Grassi,
1912). Recorded from Italy, Crimea and Transcaucasia (Börner and
Heinze, 1957).

FORDA von Heyden Pemphiginae: Fordini

An Old World genus of eight or nine species with host alternation between
Pistacia spp. (or, in one case, Toona ciliata) and roots of Gramineae. Galls
on Pistacia are formed by folding and rolling the leaf margins. The fundatrix
of most species initially forms a small temporary gall (e.g. Fig. 126F), usually
near the apex of the leaf, from which the fundatrigeniae emerge to induce the
definitive leaf-edge galls. Alatae emerging from these galls have 6-segmented
antennae, unlike those produced on the secondary hosts which always have
5-segmented antennae. The holocycle takes two years (see Fig. 3, p. 9). Anholo-
cyclic populations of several species occur outside the range of Pistacia on
roots of grasses and cereals, and several species are only known from their
secondary hosts (B & E, 1984, pp. 274-275). For accounts of the gall genera-
tions see Davatchi (1958) and Roberti (1939, 1983).

Forda formicaria von Heyden Forming half-moon-shaped ('semilunarius')

galls (Fig. 126G) on Pistacia terebinthus and P. palaestina in the Mediterra-
nean region and southwest Asia. There are also records from P. khinjuk and
P. mutica. The leaf lamina on one side is deflected down and then rolled
upwards to form a crescentic sac, yellowish when mature, the opening closed
by a curtain of fine hairs (Davatchi, 1958; Roberti, 1983). Emigrant alatae (BL
1.8-2.4 mm, with black head and thorax and green-yellow abdomen) emerge
in September-November and fly to found colonies on roots of Gramineae.
Anholocyclic populations occur on grass roots in northern Europe and North
America. Roberti (1983) gave a full description of all generations on Pistacia.
Valentyuk (1985) described a method of rearing F. formicaria on galls on
 Forda 699

P. mutica Börner (1952-3) recognized several different forms on the roots of

Gramineae in Central Europe, and sexuparae vary greatly in abdominal
chaetotaxy; it seems likely that at least three taxa are confused under the name
formicaria. 2n = 18 —23; karyotype variation was discussed by Blackman
(1987).

Forda hirsuta Mordvilko (? = mordvilkoi Börner), Plate 6e Galls are

formed by upward rolling of the leaf edge of Pistacia spp. (atlantica, khinjuk,
mutica, terebinthus, vera). They are short, sausage- or pocket-shaped lobes,
swollen and carmine red when mature, often side by side along both edges of
a leaf (Fig. 126I; Mordvilko, 1935, as follicularia; and Wertheim, 1955, as
mordvilkoi). Emigrant alatae (BL 1.5-2.3 mm) leave the galls in August-
October to colonize roots of grasses or (occasionally) bamboos (B & E, 1984,
p. 275; Halperin et al., 1989). In southwest and Central Asia; presumed
anholocyclic populations on grass roots occur in Portugal (van Harten, 1975),
and in Russia as far north as Leningrad (Mordvilko, 1935). Fet (1979) studied
the ecology of gall populations on P. vera in Turkmenia. The synonymy of
F. hirsuta is complex; if Szelegiewicz (1982) was correct in his opinion that
pawlowae Mordvilko (= dactylidis Börner) is the secondary host form of
mordvilkoi Börner, and if mordvilkoi is a synonym of hirsuta (Davatchi,
1958), then this would make the valid name for this taxon pawlowae
Mordvilko. However, the name hirsuta is retained here pending further
studies. 2n = 18 (Blackman, 1987).

Forda kaussarii Davatchi and Remaudière Galls on Pistacia khinjuk are

formed by upward folding of the leaf margin. They are flattened on the upper
side, convex with leaf veins forming slight ridges beneath; colour green becom-
ing reddish or yellowish especially on underside at maturity. Emigrant alatae
(BL 2.1-2.4 mm) leave the galls in August-October. Only the gall generations
are known, from Iran (Davatchi and Remaudière, 1957; Davatchi, 1958). This
is possibly the primary host form of F. orientalis (see below).

Forda marginata Koch Galls are formed by upward rolling of the leaf
margins of Pistacia spp. (terebinthus, palaestina) into an elongate spindle
shape (Fig. 126H), yellowish or pale green in colour. Emigrant alatae (BL
1.5-2.1mm) leave the galls in September-November to found colonies on
roots of Gramineae. Holocyclic populations occur in southern Europe,
southwest Asia and northwest India. Anholocyclic populations on grass roots
occur in northern Europe and North America (see B & E, 1984, p. 275).
Roberti (1983) fully described the gall generations and their phenology in Italy,
and Chakrabarti (1987) described the life cycle in northwest India. Karyotype
variation was studied by Blackman (1987); 2n = 17-20 (soma) or 25-40 (germ
line cells).

Forda orientalis George Aphids forming brick-red pouch galls on the

margins of the leaves of Toona ciliata in Uttar Pradesh, India were identified
as this species (A.K. Ghosh et al., 1981). However, the inhabitants of these
 700 Forda

Fig. 126. Galls of Fordini on Pistacia. A, Aploneura lentisci on P. lentiscus (after Mordvilko, 1935); B,
Asiphoniella cynodonti on P. khinjuk (after Rübsaamen, 1902); C, Baizongia pistaciae on P. terebinthus;
D, Chaetogeoica foliodentata (after Remaudière and Tao, 1958); E, Fordini 'sp. B' (final gall) on P.
atlantica; F, Forda formicaria, fundatrix gall (after Roberti, 1939); G, F. formicaria, final gall; H, F.
marginata on P. terebinthus; I, F. hirsuta on P. vera (after Mordvilko, 1935); J, F. riccobonii on
 Formosaphis 701

galls have not been described and on re-examination, A.K. Ghosh (1984b,
p. 381) considered the alatae from them to belong to a different, unidentified
Forda species. Forda kaussarii on P. khinjuk has also been suggested as the
possible primary host form of F. orientalis (Eastop and Hille Ris Lambers,
1976), which is only certainly known from grass roots in India, Pakistan and
Iran (see B & E, 1984, p. 275).

Forda riccobonii (Stephani) Pouch galls are formed on the leaves of Pistacia
atlantica; many of these may be clustered around the edge of a leaflet so that
it completely loses its shape (Fig. 126J). Emigrant alatae (BL 1.9-2.1mm)
emerge in September-November and found colonies on roots of Gramineae.
In Mediterranean area and southwest Asia. Populations on grass roots are
recorded from India (Chakrabarti et al., 1982). This species is keyed on the
basis of specimens from P. atlantica in Morocco (BMNH colln, leg. Bailie)
amd Malta (BMNH colln, leg. unknown); alatae from allegedly similar galls
in Israel (leg. E. Swirski) were much more like F. hirsuta. 2n = 18 in somatic
cells and 30 in germ line (Blackman, 1987).

Forda sichangensis Remaudière and Tao Galls on Pistacia sinensis, formed

by rolling the leaf edge upward, are elongate, crescentic in form and may
extend to the mid-rib; yellowish-green, becoming reddish-yellow when mature
(Tao, 1947; as Pemphigella marginata). Emigrant alatae, collected in May,
have blackish-brown head, thorax and appendages contrasting with yellowish-
white abdomen; BL about 2.5 mm. Their embryos have very numerous hairs
on antennae and legs (Davatchi, 1958). Other morphs unknown. Only known
definitely from China, but some emigrant alatae in the BMNH colln from galls
on P. khinjuk in Pakistan (leg. M.A. Ghani) are very similar and are possibly
this species.

FORMOSAPHIS Takahashi Pemphiginae: Fordini

One species associated with Magnoliaceae in East Asia. The alatae have
antennae with peculiar, reticulate rhinaria, and hind wings with the two oblique
veins united basally as in Pemphigini.

Formosaphis micheliae Takahashi Apterae pale brown, covered with cottony

wax; BL c. 2mm. On bark of stem and branches of Michelia spp. (alba,

P. atlantica (after Rubsaamen, 1902); K, Geoica utricularia on P. terebinthus; L, Paracletus cimiciformis on

P. terebinthus; M, Rectinasus buxtoni on P. palaestina (after Koach and Wool, 1977, as Baizongiini 'sp. B');
N, Slavum lentiscoides on P. vera (after Mordvilko, 1935); 0, S. wertheimae on P. atlantica (after Hille Ris
Lambers, 1957); P, Smynthurodes betae on P. mutica (after Mordvilko, 1929d).
702 Fullawaya

champaca, compressa, longifolia) in Japan, China and Taiwan (Takahashi,

1963b); also reported from Magnolia spp. in West Bengal (Raychaudhuri
et al., 1980e), and from Buchanania latifolia in Manipur, India (Raha et al.,
1977). The life cycle is unknown; it seems to be entirely anholocyclic. 2n = 10,
with structural heterozygosity (Blackman, 1986).

FULLAWAYA Essig Aphidinae: Pterocommatinae

Six rather large North American species living on roots of Salicaceae, closely
allied to Plocamaphis, but more hairy and with siphunculi much reduced or
absent, probably as a result of their subterranean habit. The wing venation
of the alatae seems to vary remarkably within species. Biology has been little
studied. Richards (1966b) reviewed the species (in Plocamaphis).

Fullawaya bradleyi (Richards) Apterae light brown with a pink tinge, or

flesh-coloured, with dark appendages; BL 3.7-4.3 mm. On subterranean parts
of stems of an unidentified Salix sp. in Ontario, Canada (Richards, 1966b).
Other morphs and life cycle unknown.

Fullawaya braggii (Gillette and Palmer) Apterae yellowish-brown with

blackish lateral areas or wholly dark brown to almost black, with a dusting
of wax powder along intersegmental lines; BL 2.5-3.0 mm. Alatae usually
have the media once-branched. Holocyclic on roots of an unidentified
Salix sp. in Colorado, USA (see MacGillivray, 1963). Oviparae occur in
September-November apterous males in October-November (Palmer, 1952).

Fullawaya bulbosa (Richards) Apterae light brown, pale pink or flesh-

coloured, with dark appendages; BL 3.8-4.2 mm. On subterranean parts of
stems of an unidentified Salix sp. in British Columbia, Canada (Richards,
1966b). Only apterous and alate viviparae are described; life cycle not
known.

Fullawaya ontarioensis (Richards) Colour of apterae in life unknown; BL

4.5-5.0 mm. On roots of Populus tremuloides in Ontario, Canada and also
recorded from Colorado, USA (Smith and Parron, 1978). Other morphs and
life cycle not known.

Fullawaya saliciradicis Essig Apterae have dark brown head and thorax and
yellow-brown to blackish abdomen, with a dusting of fine wax powder; BL
2.1-3.5 mm. Alatae generally have a twice-branched media. Holocyclic on
roots of Salix spp. (laevigata, longifolia). Widely distributed in North America.
Oviparae occur in October-November (Palmer, 1952). Possibly F. braggii
is a synonym, as suggested by MacGillivray (1963) and Richards (1966b).
 Geoica 703

Fullawaya terricola (Hottes and Frison) Apterae are mainly reddish-brown,

with dark appendages; BL 2.5-3.1 mm. Alatae usually have a twice-branched
media in the forewing (Hottes and Frison, 1931). On roots of unidentified
Salix in USA (Illinois, Pennsylvania, Utah). Sexuales and life cycle not known.

GEOICA Hart Pemphignae: Fordini

About nine species of Geoica are known from roots of Gramineae, and
several of these are known to host-alternate from galls on Pistacia, the
life cycle taking two years (see Fig. 3). One species (mimeuri) is monoecious
on Pistacia. The taxonomy is difficult, particularly around G. utricularia
(Brown and Blackman, 1994). Davatchi (1958) reviewed the morphs on
Pistacia.

Geoica harpazi Brown Galls on Pistacia atlantica in Israel are not clearly
identified, but probably similar in size and shape to those of G. utricularia;
they may be the P. atlantica galls described by Wertheim (1954) as green, of
rather irregular shape and with a wrinkled and uneven surface. Alate
emigrants fly in October to found colonies on roots of Gramineae
(Ammophila, Hordeum, Triticum). See Brown and Blackman (1994).

Geoica lucifuga (Zehntner) Anholocyclic populations of this species occur

on roots of Gramineae and Cyperaceae throughout the world except in cold
north temperate regions (B & E, 1984, p. 276). The holocycle has not been
described, but Mordvilko (1935) suggested that the primary host might be
Pistacia sinensis, and alatae (BL 2.1-2.2 mm) from galls on P. sinensis in
Pakistan deposited first instars with chaetotaxy agreeing well with specimens
of G. lucifuga from grass roots (G. Remaudière, pers. comm.). Very similar
alatae (BL 1.6-1.9 mm) have been collected from galls on P. khinjuk in
Pakistan (BMNH colln, leg. M. Ghani). No information is available on the
morphology of the galls in either case. 2n = 14 (from grass roots; Kulkarni,
1984).

Geoica mimeuri (Gaumont) Monoecious holocyclic on Pistacia atlantica

and (more rarely) P. terebinthus in Morocco. Galls are formed at the bases
of leaflets and resemble those of G. utricularia, but are smaller (Gaumont,
1930). The alatae leaving the galls are sexuparae; Davatchi and Remaudière
(1957) found an alata containing one embryo with normal stylets, but this
seems to be exceptional. Davatchi (1958) compared the sexuparae of
G. mimeuri with those of G. setulosa and G. utricularia.

Geoica setulosa (Passerini) Galls on Pistacia khinjuk in Iran are formed at

the bases of leaflets and are like those of G. utricularia, but larger (Davatchi,
704 Geopemphigus

1958); the gall on P. khinjuk in Israel illustrated by Koach and Wool (1977)
is probably that of G. setulosa. They open in late August-October. Emigrant
alatae (BL 1.8-2.3 mm) found colonies on roots of grasses. Anholocyclic
populations on grass roots occur in Europe (B & E, 1984, p. 276). 2n = 20
and 24 (from grass roots).

Geoica utricularia (Passerini) (= eragrostidis) This name has been applied

to a complex of species. Galls on Pistacia spp. in the Mediterranean area and
southwest Asia are roughly globular and are formed directly by the fundatrix
feeding near the base of a leaflet, close to the main vein (Fig. 126K; and
Wertheim, 1954). Colour, texture and shape seem to vary somewhat with the
host species, but on P. palaestina and P. terebinthus they are smooth and
yellowish with a pinkish tinge. Emigrant alatae (BL 1.7-2.6 mm) emerge in
July-October - the time of opening of the galls also seems to be dependent
on host species - and fly to colonize roots of Gramineae. Roberti (1983)
described the life cycle of G. utricularia sensu stricto in Italy. The considerable
variation in morphology of galls under the name G. utricularia, and of
emigrant alatae and their embryos, has led to the recognition of taxa
associated with particular Pistacia species: utricularia on P. terebinthus;
muticae Mordvilko on P. mutica; rungsi Davatchi and Remaudière on
P. atlantica; and wertheimae Brown on P. palaestina (Brown and Blackman,
1994). Koach and Wool (1977) reported electrophoretic differences between
samples from P. atlantica (perhaps G. rungsi) and P. palaestina (perhaps
G. wertheimae), indicative of discrete populations on these two primary hosts
in Israel. Wool and Koach (1976) studied the effects of genotype and environ-
ment on morphological variation in emigrant alatae of populations of this
species complex in Israel. The name utricularia is applied to anholocyclic
populations on roots of Gramineae in northern and Central Europe and
North America (B & E, 1984, p. 276), but this may not be correct. 2n = 18
(on both P. atlantica (possibly G. rungsi) and P. palaestina (possibly G.
wertheimae).

GEOPEMPHIGUS Hille Ris Lambers Pemphiginae: Fordini

Three or four species, related to Old World Baizongia and Asiphoniella, form
galls on Pistacia mexicana and P. texana in Mexico and southern USA, and
probably migrate to roots of Ipomaea spp. and various Compositae. There
is as yet no published description of the galls and gall-feeding generations,
and experimental work is required to associate the known gall-forming
and root-feeding stages. [Geopemphigus floccosus (Moreira) was described
from Ipomaea stipulata in Brazil and was subsequently found on I. batatas
and/or trapped in southern USA, Central America, the Caribbean and as
far south as Säo Paulo, Brazil (B & E, 1984, p. 279). A sexupara trapped
 Globulicaudaphis 705

in Jamaica (BMNH, leg. Coconut Industry Board) may belong to a species

found on Parthenium in Mexico and on Coreopsis in North Carolina,
USA.]

GEORGIAPHIS Maxson and Hottes Pemphiginae: Eriosomatini

Two North American species forming galls on Ulmus. Closely related to

Eriosoma, but without host alternation. The fundatrices have siphuncular
pores, and the alate sexuparae produced in the galls have the secondary
rhinaria on ANT III rather weakly developed and more widely-spaced than
in alatae of Eriosoma. Smith (1985) reviewed Georgiaphis and considered that
it should be a subgenus of Eriosoma.

Georgiaphis gillettei Maxson and Hottes Forms leaf-curl galls on Ulmus

americana and U. fulva (for illustration see Hottes and Frison, 1931, as
G. ulmi). Fundatrix and apterous fundatrigeniae greenish-blue, dusted with
wax. Monoecious holocyclic. Alate sexuparae maturing in galls in June are
greenish-brown to greenish-blue, wax-dusted; BL c. 1.5-1.6 mm (Maxson and
Hottes, 1926). In eastern North America (Illinois, Iowa, North Carolina,
Pennsylvania; BMNH colln). Synonymized with G. ulmi by Hottes and Frison
(1931), but reinstated as a distinct species by Smith (1985). Georgiaphis gillet-
tei has R IV+V equal to or longer than HT II, whereas G. ulmi has R IV+V
distinctly shorter than HT II.

Georgiaphis ulmi (Wilson) Gall is formed by curling one edge of the elm leaf
into a spindle shape, 2-5 cm long. Galled leaves turn yellow, then red, then
drop off (Wilson, 1911). Fundatrix reddish-brown. Monoecious holocyclic,
and apparently specific to Ulmus alata. Alate sexuparae, reddish-brown with
black head and thorax, are produced in second generation (Wilson, 1911),
leaving galls in May-June. In eastern North America.

GLOBULICAUDAPHIS Hille Ris Lambers

Drepanosiphinae: Phyllaphidini

One oriental species related to Myzocallis, but with a remarkably distended

CAUDA.

Globulicaudaphis pakistanica Hille Ris Lambers Apterae are 'basically pale

greenish marked with a darker olive green dorsal pattern, and with what
appear to be white paired waxy markings running laterally from the head
706 Glyphina

about two-thirds of length of body. Legs pale with black knees, antennae pale,
annulated with dark markings at joints' (Hille Ris Lambers, 1966c); BL
1.4-1.8 mm. Alatae have 2-3 rather large rounded secondary rhinaria near
base of ANT III. In colonies on undersides of leaves of Quercus dilatata in
Pakistan and India (H.P.; BMNH colln, leg. S.K. David). Biology and life
cycle unknown; apterous viviparae were collected in June, November and
December (BMNH colln). ['Myzocallis floribundi', recorded as a new species
on Q. floribunda (=dilatata) by Verma (1965) but not described, seems to
be this aphid.]

GLYPHINA Koch Thelaxinae

Five species, three palaearctic and two nearctic, associated with Alnus and
Betula. Blackman (1989) reviewed and keyed the palaearctic species, which
feed on aerial shoots, whereas the two North American species are appar-
ently both subterranean (Richards, 1968a). Parasitized by Aphidius aquilus
(= sicarius), which also attacks other Betula-feeding genera such as Betulaphis
and Calaphis (Kallistaphis spp.) (Stary, 1973).

Glyphina betulae (Kaltenbach), Plate 4c, d Apterae are dark green to almost
black with a pale spinal stripe; BL 1.2-2.0 mm. Immature stages are green.
Monoecious holocyclic, in colonies on young shoots of Betula spp., especially
B. pendula, and occasionally on Alnus spp. Ant-attended. The life cycle is
abbreviated; oviparae appear in July and apterous males in August (Heie,
1980). Not known outside Europe. 2n = 28 or 56; the significance of the
karyotype variation is uncertain (Blackman, 1989).

Glyphina longiseta Richards Apterae are reddish-brown with conspicuous

white or grey paired patches on ABD TERG 1 and 6, and dark brown to black
appendages; BL 1.9-2.1 mm. On an unidentified Alnus sp. in Nova Scotia,
Canada. The feeding site is not described but is by implication the roots or
underground stems (Richards, 1968a). Life cycle unknown.

Glyphina pseudoschrankiana Blackman Apterae are black with variable

white markings; BL 1.5-1.8 mm. Immature stages are brown. Monoecious
holocyclic, in colonies on young shoots, often on young trees, of Betula
pubescens and related downy birches, attended by ants (Blackman, 1989). In
northwest Europe and Japan. Life cycle is abbreviated, with apterous males
and oviparae occuring in July. 2n = 10.

Glyphina schrankiana Börner Apterae brown; BL 1.7-2.0 mm. Immatures

are pale brown. Monoecious holocyclic, in colonies on young growth of Alnus
spp. (incana, glutinosa), ant-attended. In Central and northern Europe. Life
 Gootiella 707

cycle is reported to be like that of G. betulae (Heie, 1980), but the sexual
morphs have apparently not been described. 2n = 8.

Glyphina setosa MacGillivray Apterae dark, reddish-brown to almost

black; BL 1.7-2.2 mm. Immature stages are light yellow-green in early
instars, tan to brownish in later instars. Ant-attended, on roots and under-
ground stems of Betula papyrifera seedlings in New Brunswick, Canada
(MacGillivray, 1963). Apterous males and oviparae were collected in August-
September.

GLYPHINAPHIS van der Goot Hormaphidinae: Cerataphidini

One species in East and Southeast Asia, characterized by lack of frontal horns
and by apterae bearing spine-like dorsal hairs, and with yeast-like extracellular
symbionts indicating an affinity with Cerataphis and Tuberaphis. Noordam
(1991) provided a generic diagnosis.

Glyphinaphis bambusae van der Goot Apterae are small pale olive green
to reddish- or greenish-brown, somewhat shiny aphids scattered on young
shoots or at leaf bases of bamboos (Bambusa spp., Phyllostachys sulphurea,
Pleioblastus latiflorus, Schizostachyum brachycladum, Shibataea kumasasa).
Immatures are spotted (Moritsu, 1983, p. 213), and alatae (BL c. 1.9 mm) have
a bluish-black abdomen (Noordam, 1991). Life cycle not recorded. In India
(Assam), China, Japan, Java and Singapore.

GOOTIELLA Tullgren Pemphiginae: Pemphigini

Two palaearctic species forming galls on Populus, related to Pachypappa but

the fundatrices have wax glands on head, thorax and abdomen, and emigrant
alatae have antennae with large star-shaped primary rhinaria. Danielsson
(1990a) reviewed the genus.

Gootiella alba Shaposhnikov Forming conical leaf-galls on Populus alba in

Russia (Uralsk), Kazakhstan and Poland. Pale yellowish-green apterae (BL
3.3-4.6 mm) live in dense wax-covered colonies on the lower side of the leaf-
cone. Alatae are unknown, suggesting monoecy (Shaposhnikov, 1952). The
uncertainties of the life cycle were discussed by Danielsson (1990a). If there
is host alternation then Aploneura juniperina Juchnevitch (1970), described
from roots of Juniperus sabina in Kazakhstan, could be the secondary host
form.
708 Greenidea

Gootiella tremulae Tullgren Forming large pale green to yellowish bag-like

galls from single leaves of Populus tremula, often situated rather high up in
old trees and therefore hard to find. The gall is formed from a longitudinally
folded leaf which is paler than the normal leaves and becomes much enlarged
with a thickened petiole (Fig. 131C, p. 803). Emigrant alatae are greyish-green,
clothed with wax, BL 3.2-4.0 mm, which in Sweden leave galls in early July
to found colonies on roots of Juniperus spp. (Danielsson, 1990b). Apterous
exules are globular, grey, covered in white wax; BL 0.9-1.5 mm. Anholocyclic
overwintering on juniper roots is common. Recorded from UK (Danielsson
and Carter, 1993b), Denmark, Fennoscandia and Poland. 2n = 16.

GREENIDEA Schouteden Greenideinae: Greenideini

About 50 East Asian species with a pattern of pale reticulation on the

siphunculi. Nearly half the species fall into subgenus Trichosiphum, which
have apterae in which this reticulation is confined to a region at the base of
each siphunculus. Biology and host relations of the genus were discussed by
A.K. Ghosh (1987). They live mainly on the shoots and young foliage of
trees, particularly Fagaceae but also Moraceae, Betulaceae, Juglandaceae and
less commonly on members of evolutionarily more recent families such as
Myrtaceae, Rosaceae and Rubiaceae. Life cycle information is only avail-
able for a few species; there seems to be much variation in the number of
parthenogenetic generations and time of appearance of sexuales. Both sexes
are winged and the eggs are somewhat flattened. Several of the most common
and widespread species seem to be mainly or entirely anholocyclic. Raychaud-
huri (1956) reviewed the world fauna and more recent accounts are (or will
be) available for Japan (Takahashi, 1962b), India (Agarwala, in press), Indian
fruit trees (A.K. Ghosh, 1975a), China (Chang and Zhong, 1979b), Taiwan
(Tao, 1962; Liao, 1978) and Java (Noordam, in prep.). At least ten aphidiid
parasitoids, including seven in or near Trioxys, and Aphelinus (Indaphelinus),
have been described from Greenidea spp. (refs: Starý and Raychaudhuri,
1982; D. Raychaudhuri, 1990).

Greenidea (Trichosiphum) anonae (Pergande) Apterae pear-shaped with

yellow-brown head and thorax and dark brown abdomen; BL 1.5-1.7 mm.
SIPH are black, curved outwards distally, about 0.2 x BL in apterae and
about 0.5 x BL in alatae. On undersides of young leaves of Annonaceae
(Anona, Artabotrys, Polyalthia) and also recorded from Antidesma, ?Bac-
cauria, Pittosporum and Symplocos. A record from Quercus sp. in India
(Chakrabarti and Raychaudhuri, 1975) needs confirmation. Sometimes ant-
attended. A single alate male is known, collected in central India in February;
probably most populations are anholocyclic. In India, Japan, Java, Malaya
and Sumatra. 2n = 22 (Khuda-Bukhsh and Kar, 1990).
 Greenidea 709

Greenidea artocarpi (Westwood) Apterae rather elongate pear-shaped, pale

brownish-green; BL 2.1-2.4 mm. SIPH are dark brown to black, long and
slender; about 0.5 x BL in apterae and 1.0 x BL in alatae. On undersides of
young leaves, usually near the main trunk, of Artocarpus spp. (heterophylla,
incisa) in southern India and Sri Lanka (David, 1956; Raychaudhuri, 1956).
Records from Southeast Asia should probably be referred to G. ficicola. Life
cycle unknown.

Greenidea bridetiae Takahashi Apterae elongate pear-shaped, yellowish-

green with dusky tips to appendages; BL 1.5-2.5 mm. SIPH are rather pale,
darker at tips, about 0.6 x BL in apterae and 0.8 x BL in alatae. On Bridelia
spp. (monoica, tomentosa) in Taiwan, Japan, Hong Kong and eastern China.
Biology unknown.

Greenidea (Trichosiphum) carpini Takahashi Apterae elongate pear-shaped,

blackish-brown with mainly black appendages; BL c. 2.1mm. SIPH about
0.25 x BL in apterae and 0.55 x BL in alate oviparae. Alate viviparae
undescribed. On Carpinus sp. in Japan. Dark brown alate oviparae and yellow
males were collected in May (Takahashi, 1963b).

Greenidea (Trichosiphum) carpinicola Banerjee and Chakrabarti Apterae

elongate oval, colour in life unrecorded: BL 1.3-1.7 mm. SIPH 0.20-
0.23 x BL in apterae and 0.50-0.54 x BL in alate oviparae. Alate viviparae
undescribed. On Carpinus spp. (faginea, viminea) in western Himalaya.
Sexuales in May (Banerjee and Chakrabarti, 1991). Possibly synonymous with
G. carpini; the small morphometric differences need to be confirmed with
similar-sized specimens.

Greenidea decaspermi Takahashi Apterae pear-shaped, yellowish-brown

with dorsal abdomen dark brown to black; BL 1.7-2.0 mm. SIPH black with
tips paler, about 0.4 x BL in apterae and 0.75 x BL in alatae (Raychaudhuri,
1956). Described from Decaspermum fruticosum in Taiwan, and David et al.
(1969) recorded it from Psidium guajava in India (although see G. neo-
ficicola). Biology is unknown.

Greenidea ficicola Takahashi Apterae pear-shaped, yellowish-brown to

dark brown; BL 1.7-2.0 mm. SIPH dark brown, curved outwards distally,
about 0.35 x BL in apterae and 0.7 x BL in alatae (Raychaudhuri, 1956). On
Ficus spp., on undersides of young leaves and on shoots, or concentrated on
the fruits. In India, Bangladesh, Nepal, eastern Siberia, China, Taiwan,
Japan, Philippines, Java, Sumatra, Malaya and Australia. Biology is little
known; anholocyclic in Taiwan (Takahashi, 1923) and no sexual morphs are
known. Sometimes ant-attended. Samanta et al. (1985) described a new
parasitoid from G. ficicola. 2n = 22.

Greenidea (Trichosiphum) flacourtiae van der Goot Apterae broadly pear-

shaped, dark brown; BL 1.4-2.4 mm. SIPH dark brown to black, 0.3-
710 Greenidea

0.4 x BL in apterae. On Distylium stellare in Java. Other morphs and biology

unknown (Raychaudhuri, 1956).

Greenidea (Trichosiphum) formosana Maki Apterae pear-shaped, dark

reddish-brown; BL 2.1-2.4 mm. SIPH yellowish-brown, darker at base and
apex, curved outwards distally, about 0.35 x BL in apterae and 0.7 x BL in
alatae. On young shoots and undersides of young leaves of Psidum guajava
and other Myrtaceae (Rhodomyrtus, Eugenia, Melaleuca). The taxonomy of
the formosana group requires clarification. Possibly G. formosana s. str.
restricts its feeding to Myrtaceae, but similar aphids occur on other plants such
as Ficus and Engelhardtia. Records of G. formosana on Ficus spp. in Taiwan
(Liao, 1978) are possibly G. fid Takahashi, described from eastern China
(Takahashi, 1937d). Greenidea guangzhouensis Chang, also described from
Ficus in China and very similar to G. formosana, may also be a synonym of
G. fid. Greenidea fici is not, however, adequately differentiated from for-
mosana ( = psidii) in Takahashi's description. Aphids identified as G. for-
mosana occur in India, Bangladesh. Nepal, eastern China, Taiwan, Japan,
Philippines, Loochoo, Java and Sumatra. Sexual morphs are unrecorded;
parthenogenetic morphs occur all year round in Taiwan (Takahashi, 1923).
Takahashi (1924a) described the immature stages. Indian populations on
guava in the BMNH collection tend to have a shorter R IV+V than those in
Southeast Asia (1.6-2.0 x HT II, as opposed to 1.9-2.2 x HT II). Greenidea
formosana-like aphids on Engelhardtia spicata and Eugenia jambolona in
India with an even shorter R IV+V (1.2-1.6 x HT II) were described as
a distinct species, G. heeri (Raychaudhuri et al., 1973). Agarwala (1983)
recorded observations of a population identified as G. formosana subspecies
heeri on guava in Bhutan. Chromosome number is 2n = 18 according to
Kulkami and Kacker (1979), but Kurl (1986) recorded chromosome numbers
of 7, 8 and 9 for various populations identified as heeri.

Greenidea (Trichosiphum) haldari Maity and Chakrabarti Apterae pear-

shaped, probably brown, with SIPH pale to brown, short, stout and out-
wardly curved; BL 2.0-2.4 mm (Maity and Chakrabarti, 1980). On an
unidentified Quercus sp. in Uttar Pradesh, India. No other morphs described,
and biology unknown.

Greenidea (Trichosiphum) kuwanai (Pergande) Apterae broadly pear-

shaped, with head, prothorax and appendages shiny mid- to dark brown; BL
2.2-2.6 mm. SIPH blackish-brown, about 0.35 x BL in apterae and up to
0.6 x BL in alatae. Immatures have a blue-grey bloom dorsally. On young
shoots and leaves of Quercus spp., with possibly valid records also from
Lithocarpus spp. and Castanea crenata; records from Castanopsis (= Shiia)
are likely to be G. okajimai and, according to Takahashi (1962b), Ray-
chaudhuri's (1956) description is partly that species. In eastern Siberia, Japan,
Korea and Taiwan. Alate oviparae occur in Japan in November-December
(Takahashi, 1923), but males have not been recorded. Greenidea bra-
chyunguis, described from Quercus sp. in Sikkim, India (Chatterjee, Mondal
 Greenidea 711

and Raychaudhuri, 1981), and G. hangnigra, described from Quercus sp. in

China (Chang and Zhong, 1979a) are very similar to, if not synonymous with,
G. kuwanai. 2n = 20 (Blackman, 1986).

Greenidea mangiferae Takahashi Apterae pear-shaped, yellowish-brown to

dark brown; BL c. 2.0mm. SIPH black, very long; almost 0.5 x BL in
apterae, and 0.75 x BL in alatae. On Mangifera indica and Euphoria longana
in Taiwan (Takahashi, 1925, 193la), and China. Biology unknown. 2n = 20
(Chen and Zhang, 1985b).

Greenidea mushana Takahashi Only the alate vivipara of this species is

described, distinguished by having more numerous secondary rhinaria on
ANT III (about 55) than any other known Greenidea species. Alatae blackish-
brown with long black SIPH, about 0.7 x BL. On young shoots of Quercus
sp. in Taiwan (Takahashi, 1925).

Greenidea (Trichosiphum) myricae Takahashi Apterae pear-shaped, green

with tips of SIPH dark brown; BL 2.1-2.5 mm. SIPH about 0.3 x BL in
apterae and 0.8 x BL in alatae. Described from specimens feeding on a young
leaf of Myrica rubra in Taiwan (Takahashi, 1925), but aphids collected two
years earlier on Alnus formosana were later identified by Takahashi as this
species (Raychaudhuri, 1956; specimens now in BMNH collection).

Greenidea neoficicola Ghosh, Ghosh and Raychaudhuri Apterae pear-

shaped, probably dark brown, with dusky yellow SIPH about 0.4 x BL; BL
c. 2.2 mm. Described from an unknown host in north Bengal (A.K. Ghosh
et al., 1970), but aphids identified as this species have subsequently been
collected on Psidium guajava in Meghalaya (Samanta et al., 1983). The latter
authors described a new braconid parasitoid genus (Indoephedrus) from this
aphid.

Greenidea (Trichosiphum) nigra Maki Apterae pear-shaped, brown-black;

BL 2.9-3.4 mm. SIPH black, about 0.3 x BL in apterae and 0.7 x BL in
alatae. On Quercus spp. and Lithocarpus rhombocarpa in Taiwan (Liao,
1978). Biology unknown. This species is not keyed as only one alate specimen
is available, and the differences between apterae of this species and those of
G. kuwanai are not clear from the literature. Takahashi (1937b) described
specimens with finer hairs as a 'variety', G. nigra kanzanensis.

Greenidea (Trichosiphum) nipponica (Suenaga) Apterae are very broadly

pear-shaped, with abdomen dorsoventrally flattened, almost circular, black
suffused with red posteriorly (Moritsu, 1983); BL c. 2.6mm. SIPH dark,
about 0.25 x BL in apterae and 0.5 x BL in alatae. On young growth of
Quercus spp. in Japan and Korea (Paik, 1972), regularly attended by ants.
Alate oviparae and males are produced in autumn and winter (Takahashi and
Sorin, 1959). 2n = 18 (Blackman, 1986). [Greenidea kumaoni, on Quercus sp.
in Uttar Pradesh, India (Chakrabarti and Raychaudhuri, 1978) is apparently
712 Greenidea

very similar and possibly synonymous, differing only in that some of the dor-
sal hairs are blunt or bifurcate.]

Greenidea (Trichosiphum) okajimai Suenaga Apterae broadly pear-shaped,

dark brown dorsally with yellow-brown head, margins of body and legs
(Moritsu, 1983); BL c. 2.0mm. SIPH blackish, about 0.34 x BL in aptera and
0.6 x BL in alata. On Castanopsis (= Shiia) cuspidata in Japan. The life cycle
is abbreviated; alate oviparae and males start appearing in early May, in the
second generation after the fundatrix (Takahashi and Sorin, 1959). Greenidea
kunmingensis, described from Castanopsis sp. in China (Chang and Zhong,
1979b), is likely to be a synonym.

Greenidea (Trichosiphum) quercicola Basu, Ghosh and Raychaudhuri

Apterae pear-shaped, colour in life unrecorded; BL c. 1.9 mm. SIPH probably
rather pale, about 0.24 x BL. On Quercus dealbata in Meghalaya, India
(R.C. Basu et al., 1975). Other morphs and biology unknown.

Greenidea quercifoliae Takahashi Apterae pear-shaped, greenish-yellow,

slightly dusky on dorsal abdomen, with legs yellowish-brown except for dark
apices (Takahashi, 1921); BL c. 1.8mm. SIPH blackish, about 0.45 x BL in
aptera and 0.73 X BL in alata. On young leaves of Quercus spp. in Taiwan;
rather uncommon, biology unknown.

Greenidea querciphaga Raychaudhuri, Ghosh, Banerjee and Ghosh Apterae

pear-shaped, colour in life unrecorded; probably rather pale with a darker
abdominal dorsal patch; BL 3.6-3.8 mm. SIPH long and pale with darker tips,
about 0.42 x BL in aptera and 0.76 x BL in alata. On an unidentified
Quercus sp. in West Bengal, India (Raychaudhuri et al., 1973). Biology and
sexuales unknown. Greenidea querciphaga is one of three Indian species
described and grouped in a separate subgenus (Neogreenidea) by Ray-
chaudhuri et al. (1973), as they all have apterae with imbrication and reticula-
tion of ANT III (and IV). The other two species, ayyari and longisetosa, were
described from unidentified hosts.

Greenidea schimae Takahashi Apterae elongate pear-shaped, green; BL

2.3-3.5 mm. SIPH long and tubular, with numerous long hairs; 0.5-0.6 x BL
and yellowish-brown in aptera, about 0.8 x BL and dark brown in alata.
CAUDA with a very distinct median process. On young leaves of Schima
spp. in Taiwan, Java (Raychaudhuri, 1956) and India (BMNH colln, leg.
K. Narayanan). Life cycle unknown.

Greenidea sutepensis Takahashi Apterae elongate pear-shaped, pale; BL

2.2-2.4 mm. SIPH pale brownish, about 0.55 x BL in aptera, alata not
known. Described from an unidentified tree, possibly a Dipterocarpus sp., in
Thailand (Takahashi, 1941; Raychaudhuri, 1956).

Greenidea symplocosis Ghosh, Basu and Raychaudhuri Apterae elongate

pear-shaped, pale (colour in life unrecorded); BL 3.1-3.3 mm. SIPH pale,
 Greenideoida 713

0.6-0.75 x BL in aptera. On Symplocos laurina in northeast India (Shillong).

Other morphs and life cycle unknown (A.K. Ghosh et al., 1969a). The dif-
ferences between this species and the preceding two described by Takahashi
(schimae, sutepensis) are not very clear.

GREENIDEOIDA van der Goot Greenideinae: Greenideini

About ten species in East and Southeast Asia described from members of
taxonomically diverse plant families. Host plants of several species, including
two from the Philippines (see Raychaudhuri, 1972), are unknown. Greeni-
deoida are rather elongate aphids distinguished from other Greenideinae by
the short last rostral segment with no clear division between R IV and R V.
The siphunculi have no reticulation and are often curved abruptly outwards
near their apices. Alatae where known are distinctive, having forewings with
an elongate, dark pterostigma and the media only once-branched, and hind-
wings much reduced without oblique veins (except in subgenera Neogreeni-
deoida and Pentatrichosiphum, which have twice-branched forewing media
and hindwing with one oblique vein). Raychaudhuri (1956) revised the species
then known, and more up-to-date accounts will soon be available.

Greenideoida ceyloniae van der Goot Apterae elongate spindle-shaped, of

variable colour, yellow, greenish-yellow or dark green (Takahashi, 1950; as
mesuae); BL 1.8-2.2 mm. Alatae are yellowish with a black head and thorax
and a large black dorsal abdominal patch. SIPH are very long, dusky to dark
but paler at apices; 0.7 x BL in aptera and 0.9 x BL in alata. On young leaves
and branches of Mesuaferrea in India, Sri Lanka and Malaya. Alate oviparae
and alate males occur in March in India (A.K. Ghosh, 1987), and fundatrices
in December (Agarwala, 1982). Agarwala and Roy (1987) studied seasonal
changes in the reproductive activity of the apterous viviparae, and Agarwala
et al. (1989b) studied developmental morphology and seasonal variation.
A.K. Ghosh et al. (1971d) described a form with shorter siphunculi from
Uvaria sp. and an unidentified plant in northeast India as a subspecies,
bhalukpongensis.

Greenideoida elongata van der Goot Apterae are very elongate-bodied,

whitish to greenish-white; BL 1.6-2.3 mm. Alatae have black head and thorax
and a large grey-black patch on the bright green abdomen. SIPH of aptera
pale brown with darker apices, 0.6-0.7 x BL; of alata 0.7-0.8 x BL. On
Bridelia tomentosa, feeding on lower sides of leaves along the mid-rib, usually
attended by ants. In Java, Malaya and Sumatra (Raychaudhuri, 1956). An
alate ovipara was collected in June in Malaya, otherwise the life cycle is
unknown. In the BMNH collection there is also another, undescribed Greeni-
deoida species from B. tomentosa in Java.
714 Grylloprociphilus

Greenideoida hannae van der Goot Apterae elongate-bodied, whitish or

pale greenish-white; BL c. 2.35 mm. SIPH rather short; about 0.3 x BL. On
young succulent shoots of Sireblus sp. in Java (van der Goot, 1917). Other
morphs and biology unknown. Raychaudhuri (1956, 1972) expressed doubts
about whether this species belongs in Greenideoida.

Greenideoida (Pentatrichosiphum) lambersi Basu Apterae elongate-bodied,

pale glassy-white; BL 2.0-2.5 mm. SIPH brown, 0.24-0.30 x BL. Found on
lower sides of leaves of Hymenodictylon sp. at an altitude of 2000m in West
Bengal, India (A.N. Basu, 1964). Raychaudhuri and Chatterjee (1977)
described the possible alata of this species. Life cycle unknown.

Greenideoida (Pentatrichosiphum) lutea Basu Apterae elongate-bodied, yel-

low; BL 2.7-3.2 mm. SIPH pale and 0.4-0.5 x BL in apterae, dark and
c. 0.55 x BL in alatae. Described from Litsea polyantha in West Bengal, India
(A.N. Basu, 1969); subsequently recorded from Cestrum fasciculatum
(A.K. Ghosh et al., 1971d) and several other plants (Raychaudhuri and
Chatterjee, 1980).

Greenideoida (Neogreenideoida) philippensis Raychaudhuri Apterae elongate-

bodied, colour in life unrecorded, probably pale; BL c. 2.3 mm. SIPH pale
and about 0.7 x BL in aptera, dark and about 0.9 x BL in alata. On
'Crotorylon celebicuni'(presumably Cratoxylon celebicum) in the Philippines
(Raychaudhuri, 1956). Biology and life cycle unknown.

GRYLLOPROCIPHILUS Smith and Pepper

Pemphiginae: Pemphigini

A genus for one North American species with distinctive wax plates in the
fundatrix, oval secondary rhinaria in alatae and first instar exules with
thickened hind femora.

Grylloprociphilus fabricator (Fitch) ( = frosti Smith and Pepper) Host-

alternating between Fagus grandifolia and roots of Taxodium distichum in
eastern USA. Fundatrices develop on undersides of leaves of Fagus, moving
to petioles and stems on maturity to form large colonies with flocculent wax.
In North Carolina, alate emigrants become adult on Fagus from April to
November. Apterous exules occur on roots of Taxodium throughout the
year, indicating partial anholocycly. Sexuparae fly from November-February
(Smith and Denmark, 1984; Denmark and Smith, 1986).
 Hamamelistes 715

HAMAMELISTES Shimer Hormaphidinae: Hormaphidini

A genus of about six species with similar host plant associations to Hor-
maphis, but with a gall (or 'pseudogall') dwelling stage on Betula as well as,
or instead of, Hamamelis. Alatae have two oblique veins in the hindwing.
Species-level taxonomy is problematic and needs more biological studies for
clarification.

Hamamelistes betulinus Horvath, Plate 3c, d Apterae in pseudogalls on

birch are dark brown to black; BL about 1.5mm, secreting white wax. They
have very short, 3-segmented antennae and no siphunculi. Alatae (BL 1.3-
2.0mm) have 5-segmented antennae and pigmented siphuncular pores.
Hamamelistes betulinus is anholocyclic on Betula spp., mainly on B. pendula,
in Europe and northern Asia, east to Mongolia and China. It feeds on the
undersides of the leaves causing pale yellowish blisters to develop on the upper
surfaces (Fig. 127), and overwinters as aleyrodiform/coccidiform first instar
larvae on the twigs. For a more detailed account see Heie (1980). A 'sub-
species', makabiae Inouye, has been described from B. maximowicziana in
Japan. Hamamelistes shirakabae Monzen, on B. japonica in Japan, appears
from the description to be very similar to H. betulinus, except that its alatae
are reported to lack, or have indistinct, siphunculi. 2n = 12.

Fig. 127. Hamamelistes betulinus on Betula leaves, showing blisters on upper surface and aphids
feeding beneath (from Mordvilko, 1935).
716 Hamamelistes

Hamamelistes cristafoliae Monzen Apterae are reddish-yellow and inhabit

cockscomb-like galls projecting from the upper surfaces of leaves of B. maxi-
mowicziana in Japan (Monzen, 1954). The life cycle is not known.

Hamamelistes gibberi Monzen Greenish apterae with dark brown legs

inhabit pseudogalls on the undersides of leaves of B. ermanii in Japan, the
upper surfaces blistering and turning yellowish in response to feeding. Apterae
differ from those of H. betulinus (and H. shirakabae) in having 4-segmented
antennae and siphuncular pores. 2n = 12 (Blackman, 1986; wrongly identified
as H. betulinus). Similar aphids feed on B. grossa var. ulmifoliae (H. gibberi
'biological race grossae'). See Monzen (1954) for more detail.

Hamamelistes kagamii (Monzen) Galls on flowerbuds of Hamamelis

japonica in Japan are large, stalked, pouch-like, with 7-13 horn-like pro-
tuberances, yellowish and covered with brownish hairs (Monzen, 1954). Apart
from its gall this aphid strongly resembles H. miyabei.

Hamamelistes miyabei (Matsumura) This aphid produces rather large

yellowish-green spiny galls on the buds of Hamamelis japonica in Japan,
similar to those of H. spinosus in North America. Hamamelistes miyabei does
not migrate to Betula in Japan according to Monzen (1954), but Bhattacharya
et al. (1983) found a similar or identical aphid causing folded-leaf pseudogalls
on Betula alnoides in northern India. (They described an apterous ovipara
from these galls, perhaps indicating a distinct species.)

Hamamelistes spinosus Shimer Apterous fundatrices in the spiny galls on

Hamamelis virginiana (Fig. 128A) are about 2 mm long, almost globular, dark

Fig. 128. Galls of Hormaphidini on Hamamelis: A, Hamamelistes spinosus and B, Hormaphis

hamamelidis (both from Pergande, 1901).
 Hannabura 717

purplish-brown, covered with wax meal. Apterae feeding on the undersides of

birch leaves are smaller, dark brownish-red to purple with abdominal tufts of
white wax. There is also a black, rugose, coccid-like morph that overwinters
on birch twigs. The alatae may be distinguished from those of Hormaphis
hamamelidis migrating between the same hosts by their 5-segmented antennae
and two oblique veins in the hindwing. This species occurs throughout North
America. The life cycle is complex and extends over two years, as determined
by Eergande (1901). Large numbers of alatae emerge from the galls founded
by fundatrices in spring on H. virginiana and fly to Betula spp. (papyrifera,
lutea, nigra), where an overwintering coccidiform morph lives on the twigs
until the following spring. The progeny of this morph move to the undersides
of young leaves, producing reddish-brown to crimson-coloured swellings of
the leaf lamina between the veins. The return migrants to witch hazel produce
sexuales which lay overwintering eggs. 2n = c. 50*.

HAMILTONAPHIS Aoki, Kurosu and Fukatsu

Hormaphidinae: Cerataphidini

A genus for one species monoecious on Styrax, related to Cerataphis and

having similar yeast-like extracellular symbionts (Aoki et al., 1993).

Hamiltonaphis styraci (Matsumura) Monoecious, forming large, coral-like

galls on the twigs of Styrax obassia in Japan. This is the only known
monoecious gall-former on Styrax. The galls take two years to develop (Aoki
and Kurosu, 1990; as Astegopteryx). Alate sexuparae (BL c. 1.4mm) appear
in the August of the second year and deposit yellowish sexuales on the under-
sides of the Styrax leaves. The sterile second instar soldiers in the gall not only
defend their colony but also clean the gall, by pushing out honeydew droplets
and exuviae (Aoki and Kurosu, 1989; as Astegopteryx).

HANNABURA Matsumura Drepanosiphinae: Phyllaphidini

A genus for one East Asian species related to Calaphis but with fewer hairs
on the first tarsal segment.

Hannabura alnicola Matsumura Apterae pale yellowish, with mainly whitish

appendages (Matsumura, 1917); BL 1.5-1.6 mm. On Alnus spp. (hirsuta
var. sibirica, japonica, matusumurae) in Japan. Oviparae were collected in
July-August in Hokkaido (Higuchi, 1972); biology otherwise unknown.
718 Heminipponaphis

HEMINIPPONAPHIS Chakrabarti and Raha

Hormaphidinae: Nipponaphidini

One species, H. querciphaga, is described, from Quercus incana in northern

India (Chakrabarti and Raha, 1985). This seems likely to be a synonym of
Pseudothoracaphis himachali, q.v.

HEMIPODAPHIS David, Narayanan and Rajasingh

Pemphiginae: Eriosomatini

Two species, one known only from galls on Zelkova in Japan and the
other known only from a monocotyledenous secondary host in India. The
taxonomie position of the genus was reviewed by Akimoto (1983).
Hemipodaphis persimilis Akimoto Forming leaf-roll galls on Zelkova ser-
rata. Emigrant alatae are very small (BL 0.6-0.7 mm), brown, emerging in
July-August to migrate to an unknown secondary host (Akimoto, 1983).
Populations persist in the galls until October (Akimoto, 1992). Aggressive
behaviour by first instar larvae was described by Aoki (1978; as Colopha
sp. nr caucasica). 2n = 36 (Blackman, 1986).

HOPLOCALLIS Pintera Drepanosiphinae: Phyllaphidini

Four palaearctic species related to Myzocallis, but with processus terminalis

always shorter than ANT VI base, with small pronotal hairs in clusters and
with distinctive dorsal pigmentation and sclerotization. All viviparae are alate.
Immatures are much broader and flatter than adult alatae and have 4 longi-
tudinal rows of brown sclerites bearing clusters of hairs. Quednau and
Barbagallo (1991) keyed the species.
Hoplocallis microsetosus Quednau and Chakrabarti Colour in life not
recorded; BL of alata c. 2.4 mm. On Quercus incana in Uttar Pradesh, India
(Quednau and Chakrabarti, 1976) and also on Q. baloot (= Q. ilex) in
Afghanistan (BMNH colln, leg. DHRL). Sexuales unknown.
Hoplocallis microsiphon Quednau and Barbagallo Colour in life not
recorded; BL of alata 1.6-2.1mm. On various Quercus spp., but perhaps
 Hoplochaetaphis 719

usually on Q. calliprinos, in the Mediterranean region (Israel, Lebanon, Sicily,

Turkey, Greece) and former Czechoslovakia (Quednau and Barbagallo, 1991).
Life cycle unknown; an alate male was collected in Greece in December.

Hoplocallis pictus (Ferrari) Alatae rather elongate, pale yellow-green with

banded antennae and a series of dark spinal markings; BL 1.8-2.2 mm. On
undersides of leaves of Quercus spp. throughout the Mediterranean area,
in southwest Asia, and introduced into South Africa and South America
(Argentina, Chile). Monoecious holocyclic; in Israel males and oviparae
appear in December (Swirski, 1953; as Myzocallis bodenheimeri). 2n = 14.

Hoplocallis ruperti Pintera Alatae rather elongate, yellow to whitish with

dark lateral as well as spinal markings; BL c. 2.0mm. Immatures are yellow
with contrasting dark brown quadrate dorsal patches. On leaves of Quercus
spp. (cerris, robur) in Central and southeast Europe (Austria, Bulgaria,
former Czechoslovakia, Hungary, Turkey). Monoecious holocyclic; oviparae
and males are produced in October in former Czechoslovakia (Pintera, 1952).

HOPLOCHAETAPHIS Aizenberg Drepanosiphinae: Phyllaphidini

A genus for one small oak-feeding species in southern Europe and southwest
Asia. Colonies include apterous viviparae with very long dorsal hairs; they
appear to be adapted to living in leaves rolled by caterpillars.

Hoplochaetaphis zachvatkini (Aizenberg and Moravskaya) Apterae are

broadly pear-shaped, pinkish-red, rich red, dull brown or orange, with a
variable blackish-brown dorsal sclerotic pattern; sometimes covered with a
thin mealy wax so that they resemble caterpillar faeces (Aizenberg and
Moravskaya, 1959); BL 0.8-1.1 mm. Immatures are reddish. Alatae have dark
head and thorax and paired dark patches or cross-bands on ABD TERG 3-7
or 4-7, or a dark patch centred on ABD TERG 4-5, and a conspicuous dark
patch at the distal end of Cu lb in the forewing. On Quercus spp., often in
leaves rolled by caterpillars. In the Mediterranean area and southwest Asia.
Monoecious holocyclic; sexual morphs were recorded in October in Turkey
(on Quercus sp.; Tuatay and Remaudière, 1964), Iran (on Q. persica; BMNH
colln, leg. S. Hodjat) and Italy (on Q. pubescens; Patti, 1983, as H. parvula).
Vereshchagin and Aukshtikal'nene (1981) made ecological observations.
[H. parvula Hille Ris Lambers and van den Bosch is a synonym (W. Quednau,
pers. comm.); there is considerable seasonal variation in extent of dorsal
sclerotization and form of dorsal hairs of this species.]
720 Hoplochaitophorus

HOPLOCHAITOPHORUS Granovsky
Drepanosiphinae: Phyllaphidini

Three North American oak-feeding species apparently related to Neo-

symydobius. Colonies consist mainly of apterae, which typically have
Chaitophorus-like pigmented spiny dorsal hairs.

Hoplochaitophorus heterotrichus Quednau 'Alatiform apterae' whitish-

green with banded antennae, a chestnut brown spot covering the middle of
the thorax and a pattern of dorsal abdominal markings (Quednau, 1971); BL
c. 2.2mm. Alatae have dark-bordered forewing veins. On basal part of lower
leaf surface of Quercus prinus in northeastern USA (Pennsylvania, New
York). This species is possibly synonymous with H. quercicola (see below).

Hoplochaitophorus quercicola (Monell) Apterae greenish with dark dorsal

markings; BL 1.8-2.1mm. Alatae with dark-bordered forewing veins. First
described from specimens colonizing undersides of leaves of Quercus prinus
in Illinois, USA (Monell, 1879), but Granovsky (1933) recorded it living in
dense colonies along veins on upper surface of foliage, and according to
Thomas (1879; as Callipterus quercifolii) it occurs on both sides of leaves
along the veins.* Recorded from various Quercus spp. (alba, macrocarpa,
prinus, rubra, etc.) in North America (except the northwest). Monoecious
holocyclic: apterous males and oviparae (the latter with a long, drawn-out
ovipositor) occur in October (in Illinois).

Hoplochaitophorus spiniferus Quednau Colour in life not recorded, hind

tibiae of apterae dark brown; BL of aptera c. 2.3 mm. On Quercus bicolor
in New York, USA (Quednau, 1972). Biology and other morphs unknown.

HOPLOTHORACAPHIS Pramanick,
Samanta and Raychaudhuri Hormaphidinae: Cerataphidini

One Indian species resembling Parathoracaphis, but without dorsal reticula-

tion and with normally developed tarsi and claws.

* [Quednau (1971) recorded H. quercicola as feeding specifically on upper sides of leaves of 0. macrocarpa, but this is
clearly not the case. In fact Monell (1879) described his aphid from the same host and feeding site as Quednau's
heterotrichus. The latter was described from individuals intermediate between apterae and alatae, and further work may
reveal it to be a synonym of quercicola; besides varying seasonally, apterous specimens with different degrees of alatiformity
may have varying proportions of spine-like and finer dorsal hairs and also differ correspondingly in the length and form
of antennal hairs.]
 Hormaphis 721

Hoplothoracaphis manipurensis Pramanick, Samanta and Raychaudhuri

Apterae aleyrodiform, brown, oval; BL c. 0.9mm. On Litsea sebifera in
Manipour, India (Pramanick et al., 1983). Biology unknown. (An apterous
morph collected on Amaranthus in another locality was described as the para-
type fundatrix of this species, but no life cycle observations or other evidence
linking these two morphs was reported, and Amaranthus is a most unlikely
primary host for a hormaphidine.)

HORMAPHIS Osten-Sacken Hormaphidinae: Hormaphidini

A genus for two or three species migrating from Hamamelis to Betula, or

monoecious on Hamamelis. Conical pouch galls protruding from the upper
sides of the leaves are produced on Hamamelis, but the apterous forms on
Betula are aleyrodiform with a fringe of white wax, living freely on the leaves.
The alatae have 3- or 5-segmented antennae and a single oblique vein in the
hindwing.

Hormaphis betulae (Mordvilko) Plate 3b Small, flattened, subcircular

yellowish-green or yellowish-brown aphids with a fringe of radiating wax
filaments, dispersed on undersides of birch leaves, not causing any leaf defor-
mation. Legs and antennae are greatly reduced. Alatae produced on birch
differ from those of the other European birch-feeding hormaphidine,
Hamamelistes betulinus, in having no siphunculi and usually only one oblique
hindwing vein. Hormaphis betulae is anholocyclic on Betula spp. (pendula,
pubescens, nana) in northern and Central Europe and in Siberia. It is believed
that overwintering may occur as a larval stage in the soil, and that the
aleyrodiform generations may have only three larval instars (see Heie, 1980).
Recently it was shown that a holocycle exists in Japan and that the aphid
described as H. gallifoliae (Monzen, 1954), forming conical pouch galls on the
upper sides of leaves of Hamamelis japonica, is the primary host form of
H. betulae (Aoki and Kurosu, 1991c). Gall-cleaning behaviour was studied by
Kurosu and Aoki (1991a). Alatae leave the galls in May-June to deposit larvae
on leaves of Betula platyphylla var. japonica. 2n = 18 (Blackman, 1986).

Hormaphis hamamelidis (Fitch) Apterous fundatrices in conical galls on

Hamamelis (Fig. 128B, p. 716) are dark purple with white wax on the
abdomen. Apterae on birch leaves are very small, dark brown to black, with
a fringe of radiating white wax filaments. Alatae have only three antennal
segments, one oblique vein in the hindwing, and no siphunculi. Two types of
life cycle occur in eastern North America. At low elevations and latitudes
south of 41 °N alatae migrate from the galls on Hamamelis virginiana and have
three aleyrodiform generations on Betula nigra, whereas on high ground or
more northerly latitudes there is an abbreviated life cycle on witch hazel
722 Hyalomyzus

without host alternation (von Dohlen and Gill, 1989). Both life cycles occur
together at intermediate locations, so it is likely that the same name is being
used for two sibling species. 2n = 18 (for both forms)*.

HYALOMYZUS Richards Aphidinae: Macrosiphini

A Myzus-like genus with seven North American and three Indian species,
related to Ovatus but with swollen siphunculi and with a strongly wrinkled
tergum in apterae. Three American species apparently host-alternate between
woody Rosaceae (usually Crataegus) and secondary host plants in several
families; however, as pointed out by Voegtlin (1984), there is in no case any
published evidence of host transfer tests to verify the life cycle. The other
species are known only from herbaceous plants. Smith (1982) keyed the North
American species and Voegtlin (1984) reviewed their biologies. L.K. Ghosh
(1986) reviewed the species in Himachal Pradesh, India.

Hyalomyzus eriobotryae (Tissot) Apterae brown; BL c. 1.8mm. Alatae are

Myzus-like but have secondary rhinaria on ANT III, IV and V and no dark
dorsal abdominal patch. On young growth of Crataegus spp. in spring in
eastern North America (Ontario to Florida) and also recorded from North
Dakota and Washington. Originally described from alate ?gynoparae and
males collected in December in Florida on Eriobotrya japonica, and at the
same time recorded from apple and hawthorn (Tissot, 1935). Crataegus spp.
are presumably the normal native primary hosts. Apparently heteroecious
holocyclic, with the presumed secondary host populations recorded from
Labiatae (Lycopus virginicus and Collinsonia canadensis).

Hyalomyzus sensoriatus (Mason) Only alatae (?fundatrigeniae) were

described from the primary host, Crataegus crusgalli, in Washington, DC
(Mason, 1940). The secondary host is purportedly Hypericum sp., based on
collections in Pennsylvania (Nielsson and Habeck, 1971), but alate males and
gynoparae collected on Lycopus americana on Long Island, New York
(BMNH colln, leg. M.D. Leonard) also agree well with sensoriatus, so the life
cycle requires confirmation. Also recorded from Quebec, Canada.

Hyalomyzus tissoti Nielsson and Habeck The only morph described from
Crataegus spp. (praeformosa, vicana) is the alate ?fundatrigenia; it has brown
thorax and yellowish-brown to brown abdomen; BL 1.0-1.5 mm (Nielsson and
Habeck, 1971). 'Secondary hosts' are recorded as Drosera spp. and Isnardia
intermedia, but the species is only known from Florida, where anholocyclic
populations are likely, so the host alternation requires confirmation.
 Hydronaphis 723

HYALOPTERUS Koch Aphidinae: Aphidini

A genus once used in a much broader sense to include most Aphidinae with
small siphunculi, but now restricted to two or three very similar species of
elongate-bodied, mealy aphids that alternate between Prunus spp. and
Phragmites. Stroyan (1984) and Heie (1986) reviewed the genus; see also B &
E, 1984, p. 282-283. Basky and Szalay-Marszo (1987) studied morphological
differences between H. amygdali (from peach) and H. pruni (from plum). The
couplet provided in the Prunus key takes account of their work, but will
still be unreliable for single specimens. Spampinato et al. (1988) studied
electrophoretic differences which indicate that 'H. amygdali probably includes
two sibling species, one mainly colonizing P. amygdalus and the other favour-
ing P. persica as primary host. Basky (1982) studied natural enemies and
Huang et al. (1986) studied phenology of H. amygdali in China. 2n = 10
(for pruni).

HYDRONAPHIS Miyazaki Aphidinae: Macrosiphini

Five Myzus-like oriental species, resembling Cryptaphis in having long hairs

and a sclerotized tergum, but the hairs are not apically expanded and the
siphunculi are slightly clavate. One species alternates between Corylopsis and
the roots of Impatiens in Japan, while the others are recorded from various
presumed secondary host plants, usually feeding on the roots (Miyazaki,
1971).

Hydronaphis impatiens Shinji In early-spring colonies living on undersides

of leaves of Corylopsis spp., without causing any leaf deformation. Apterae
(fundatrices) are pale yellow, with dark appendages and black-tipped siphun-
culi; BL c. 2.4 mm. Progeny of fundatrices are all alate, migrating in
April-May to found colonies on roots and subterranean stems of Impatiens
spp.; the emigrant alatae have a yellow abdomen with a black dorsal patch.
Alate males and gynoparae return to Corylopsis in October (S. Takahashi,
1986). Only known from Japan.

Hydronaphis laporteae Miyazaki Apterae broadly oval, dull yellow with

antennae, legs and siphunculi mainly pale but dark at apices; BL 1.6-1.8 mm.
Described from specimens collected on undersides of leaves of Laportea bulbi-
fera (Miyazaki, 1971), but apparently the normal feeding site is the roots
(S. Takahashi, 1986). Only recorded from Japan and the life cycle is unknown.
lít!

724 Hysteroneura

HYSTERONEURA Davis Aphidinae: Aphidini

One originally North American species in which the CAUDA is long and pale
and the hindwing has only one oblique vein.

Hysteroneura setariae (Thomas) Apterae are dark rusty brown with dark
siphunculi, contrastingly pale sections of the antennae and tibiae and a
pale CAUDA; BL 1.3-2.3 mm. In spring on twigs and leaves of tender new
shoots of Prunus domestica in North America. Host alternation occurs
to aerial parts of Gramineae, but ant-attended populations may persist on
plum throughout the year (Gillette and Taylor, 1908). Anholoyclic popula-
tions of H. setariae have become widely distributed throughout the warmer
regions of the world on Gramineae (see B & E, 1984, pp. 285-286), including
bamboos. Carver (1976) found sexuales and eggs on Prunus persica in
southern Australia, but it is not known if the holocycle was completed.
2n = 12.

ILLINOIA Wilson Aphidinae: Macrosiphini

About 45 mostly North American species, recognized most easily by the

weakly to moderately clavate siphunculi, similar to those of Amphorophora
but ornamented on the distal, constricted region with a few rows of poly-
gonal reticulation. Evolution of host relationships within the genus is some-
what obscure, but may be connected with the ability to feed on plants
protected from other small insects by sticky hairs; many species in the
two principal subgeneric groups are associated with Ericaceae, but others
feed on taxonomically diverse plants (B & E, 1984, p. 286), including trees.
MacGillivray (1958) revised the genus Illinoia under the name Masonaphis.

Illinoia alni (Mason) Apterae spindle-shaped, pale green with a dark green
median stripe and dark spots at bases of siphunculi; BL 2.5-3.1mm.
Immatures pale green. In small colonies on undersides of leaves of Alnus
rugosa in northeastern USA and eastern Canada (MacGillivray, 1958).
Monoecious holocyclic. 2n = 10.

Illinoia azaleae (Mason) Apterae green; BL 1.9-2.2 mm. Immatures pale

greyish-green. Mostly on Rhododendron spp. (especially azaleas) and other
Ericaceae, but sometimes collected (in North America) on Myrica asplenifolia
which apparently can function as a reserve host (MacGillivray, 1958; Stroyan,
1972). Probably mainly anholocyclic; oviparae sometimes occur, but males are
 Illinoia 725

undescribed. Introduced from North America into Europe, New Zealand and
South Africa. 2n = 10.

Illinoia borealis (Mason) Colour in life unknown, probably dark green or

brownish-green with dark siphunculi; BL 1.7-2.5 mm. Described from Gaul-
theria procumbens, but also collected from Vaccinium sp. and Myrica
asplenifolia (leaves) (MacGillivray, 1958). Only known from Maine, USA.
Life cycle unknown.

Illinoia canadensis (MacGillivray) Apterae rather shiny, yellowish-green

mid-dorsally, darker green laterally with dark spots at the bases of the dark-
tipped siphunculi; BL c. 2.7 mm. On Myrica sp. in New Brunswick, Canada
(MacGillivray, 1958). Life cycle unknown.

Illinoia corylina (Davidson) Described from immature and adult alatae on

Corylus rostrata (= cornuta var. californica), but that is the only record from
Corylus, it being more commonly found on Physocarpus capitatus (Mason,
1925, quoting Davidson). Apterae are undescribed (MacGillivray, 1958). In
California, USA. Life cycle unknown.

Illinoia lambersi (MacGillivray) Apterae and alatae with three distinct

colour morphs, green, pink and yellow, which can occur in mixed populations;
BL 2.0-3.7 mm. The principal host is Rhododendron, but colonies sometimes
occur on Ilex aquilifolium (British Columbia, Canada, Forbes and Chan,
1989; and England, BMNH colln, leg. VFE). In western North America and
introduced into Europe and Chile (BMNH colln, leg. R. Carrillo). Holocyclic
on Rhododendron in California, USA, but European populations are anholo-
cyclic (Hille Ris Lambers, 1973a). 2n = 10.

Illinoia liriodendri (Monell) Apterae spindle-shaped, pale green lightly

dusted with wax, with antennae and siphunculi black except at bases and legs
pale green except for black tibial apices and tarsi; BL 1.7-2.5 mm. A red
colour form also occurs. Monoecious holocyclic on Liriodendron tulipifera in
North America, feeding on undersides of leaves. Oviparae and alate males in
October. Davis (1909) gave a full description, Dreistadt (1987) studied honey-
dew production and Dreistadt and Dahlsten (1988) discussed the timing of
control measures. 2n = 10.

Illinoia morrisoni (Swain), Plate 14e, f Apterae spindle-shaped, long-legged,

rather dark apple green, with antennae dark beyond the base of ANT III and
legs and siphunculi tipped with black; BL 1.5-2.3 mm. On terminal leaves and
shoots of numerous conifers, usually Cupressaceae (Actinostrobus, Callitris,
Calocedrus, Chamaecyparis, Cupressocyparis, Cupressus, Juniperus, Thuja,
Widdringtonia); also on Araucariaceae (Araucaria) and Taxodiaceae (Metase-
quoia, Sequoia, Taxodium). There is only one record from Pinaceae (Cedrus
deodora; BMNH colln, leg. H.G. Walker). In western North America and
introduced into England, but apparently it has not spread more widely. Life
726 Indiochaitophorus

cycle unknown; no sexual morphs are described and, in California at least,

it seems to be anholocyclic. 2n = 10.

Illinoia patriciae (Robinson) Apterae spindle-shaped, yellowish-green to

green; BL 1.7-2.3 mm. On new needles of Tsuga heterophylla in British
Columbia, Canada (Robinson, 1969). Differences from the one other conifer-
feeding Illinoia species, I. morrisoni (which has not been recorded from
Tsuga) are given by Robinson. Life cycle not known.

Illinoia wilhelminae (Hille Ris Lambers) Apterae spindle-shaped, colour in

life not recorded; BL c. 3.1 mm. The siphunculi show little or no trace of swell-
ing. Described from a single aptera collected in Quebec, Canada, on Alnus
rugosa, but the form of R IV+V suggests that this was not its normal host
plant (Hille Ris Lambers, 1962b). Not recorded since.

INDIOCHAITOPHORUS Verma
Drepanosiphinae: Phyllaphidini

One elm-feeding species related to Sinochaitophorus, but the aptera has short
antennal hairs and a different pattern of dorsal sclerotization.

Indiochaitophorus furcatus Verma Apterae rather elongate oval, pale

brown with thick black dorsal longitudinal stripes; BL 1.3-2.0 mm. Alatae
have paired dark spinal and marginal sclerites on each of ABD TERG 1-6.
Recorded from undersides of leaves and twigs of Ulmus wallichiana in
Kashmir, India and on U. laevigata in Pakistan (BMNH colln, leg. M.A.
Ghani). Heavy infestation of riverside trees was reported in Kashmir in June
(Verma, 1970a). Life cycle unknown.

INDONIPPONAPHIS Ghosh and Raychaudhuri

Hormaphidinae: Nipponaphidini

Two little-known species described from Quercus, related to Nipponaphis but

without spinal hairs on ABD TERG 7, and with paired spinal and marginal
tubercles on prosoma and abdominal plate.

Indonipponaphis fulvicola Sorin Apterae brown, almost circular, dorso-

ventrally flattened, strongly sclerotized; BL c. 2.1mm. On branches of
Quercus acuta in Japan (Sorin, 1979b). Biology unknown.
 Kaburagia 727

Indonipponaphis tuberculata Ghosh and Raychaudhuri Apterae brownish,

oval, strongly sclerotized; BL 1.9-2.1mm. On stem of Quercus dealbata in
Meghalaya, India (A.K. Ghosh and Raychaudhuri, 1973a). Biology and other
morphs unknown.

INDOREGMA Chakrabarti and Maity

Hormaphidinae: Cerataphidini

One species in India, closely related to Ceratovacuna.

Indoregma bambusae Chakrabarti and Maity Appearance in life not

recorded, aptera probably rather pale, with wax; BL 2.0-2.7 mm. On Bam-
busa sp. in Uttar Pradesh, India (Chakrabarti and Maity, 1982). Biology
unknown. Perhaps a synonym of Ceratovacuna indica, q.v.

INDOTUBEROAPHIS Chakrabarti and Maity

Aphidinae: Macrosiphini

One species on Sorbus in India, with many distinctive features including

paired spinal processes on ABD TERG 1-4 but only single median processes
on ABD TERG 5-8.

Indotuberoaphis sorbi Chakrabarti and Maity Apterae pale brown (?);

BL 2.5-2.7 mm. On undersides of young leaves of Sorbus foliolosa in
Uttar Pradesh, India (Chakrabarti and Maity, 1984). Biology and other
morphs unknown. Later generations may have smaller abdominal processes.

KABURAGIA Takahashi Pemphiginae: Fordini

Three or more oriental species related to Melaphis, Nurudea and Schlechten-

dalia, but the alatae from galls on Rhus have distinctive antennal sensoriation.
A.K. Ghosh (1984b) reviewed the genus and Xiang (1980/81) reviewed the
species in China.
728 Kaltenbachiella

Kaburagia ailanthi Chowdhuri, Basu, Chakrabarti and Raychaudhuri Emi-

grant alatae from galls are rather elongate-bodied, colour in life unrecorded:
BL 1.7-1.8 mm. The host plant given in the original description, and on
the paratype slides in the BMNH collection, is Ailanthus sp., without any
mention of gall formation (Chowdhuri et al., 1969). Chakrabarti et al.
(1985) reported that it forms a petiole gall on Ailanthus glandulosa, but no
new collection data were given in that paper and the unusual host association
requires additional confirmation. A more probable host plant would be
Rhus punjabensis. In India (Himachal Pradesh, Uttar Pradesh). Life cycle
unknown.

Kaburagia ovatirhusicola Xiang Galls on leaflets of Rhus potaninii in China

(Shansi) are obovate, green, with prominent net-like veins on surface. Differ-
ences from K. ovigallis (= rhusicola?) in the morphology of the emigrant alata
(BL 1.5-2.0 mm) were described by Xiang (1980/81). Secondary host is
unknown.

Kaburagia rhusicola Takagi Galls on leaflets of Rhus (potaninii, punja-

bensis var. sinica) are elongate pear- or fig-shaped, somewhat pointed at distal
end, 3-10 cm long, with some longitudinal striations, yellowish-green when
mature (Takagi, 1937; Xiang, 1980/81). Alate emigrants are dark grey to black
(presumably with wax secretion); BL 1.3-1.5 mm. Emigration from galls in
northern China occurs in late summer, hibernation occurs as immature stages
on the secondary host, the moss Erythrodontium leptothallum, and sexuparae
return to Rhus in early spring (Xiang, 1980/81). In China and Korea (Chosen).
Macrorhinarium ensigallis and M. ovogallis Tsai and Tang have been
synonymized with K. rhusicola (Eastop and Hille Ris Lambers, 1976), although
Xiang (1980/81) treated both these as distinct taxa.

KALTENBACHIELLA Schouteden Pemphiginae: Eriosomatini

Eight species primarily associated with Ulmus and related to Colopha, but the
hindwing has two oblique veins, and the feeding position of the first instar
fundatrices is such that the gall arises from the mid-rib of the leaf near its
base, rather than from the leaf lamina. A.K. Ghosh (1981) reviewed the genus,
and Akimoto (1985a) reviewed the species in East Asia.

Kaltenbachiella carpinicola Ghosh, Chakrabarti and Bhattacharya Reported

to form an open gall consisting of a swollen cup-shaped pouch or cavity on
leaf blades of an unidentified Carpinus sp. (A.K. Ghosh et al., 1981). Alatae
collected from these galls in June have an unbranched forewing media and
secondary rhinaria distributed ANT III 20-29, IV 7-10, V 10-13 and VI 15-17;
BL 2.2-2.6 mm. Life cycle is unknown. The host plant and position of the
 Kaltenbachiella 729

gall are anomalous and need additional confirmation, being based on only one
collection. In India (Uttar Pradesh).

Kaltenbachiella glabra Akimoto Galls are almost globular, unstalked,

smooth, projecting upward from the mid-rib of the leaf. Only the gall and
fundatrix are described (Akimoto, 1985a), so this species could not be included
in the key to aphids on Ulmus. On Ulmus uyematsui in Taiwan. Life cycle
is unknown.

Kaltenbachiella japonica (Matsumura) Galls are globular, covered with fine

spine-like projections, green, projecting upward from the mid-rib, which
tends to be bent downwards where the gall is attached (Fig. 134K, p. 898).
Monoecious holocyclic on Ulmus spp. On U. japonica in Japan the galls occur
mainly on mature trees, especially on shoots growing directly from the trunk
and may occur four or more to a leaf. Alate sexuparae (BL 1.4-1.7 mm)
emerge from galls in July-August and produce sexuales in crevices of the
trunk, often near their old galls (Akimoto, 1985a). In Japan and Siberia.
[Alate specimens trapped in Sri Lanka (BMNH colln, leg. E. Judenko) and
described as K. japonica by A.K. Ghosh (1984b), have embryos with mouth-
parts and are closer to K. nirecola, except that they lack siphunculi.] Evolution
of the monoecious life cycle of K. japonica was discussed by Akimoto (1985c).
2n = 16 (Blackman, 1986).

Kaltenbachiella nirecola (Matsumura) Galls on Ulmus spp. are oblong, bag-

shaped, with a narrower section at the base, projecting upward from the mid-
rib of the leaf; green, often with reddish tinge distally, covered in short whitish
hairs. Emigrant alatae have the forewing media usually once-branched but
sometimes unbranched; BL 1.3-1.7 mm. In Japan they emerge in July and
found colonies on the roots of Polygonum Orientale (Akimoto, 1985a). In
Japan and Siberia.

Kaltenbachiella pallida (Haliday) Galls on Ulmus spp. are globular, pale,

densely covered in short fine hairs, projecting from the mid-rib mainly on the
upper side of the leaf (Fig. 134H, p. 898). Emigrant alatae have the forewing
media usually unbranched, sometimes once-branched; BL 1.8-2.1 mm. They
emerge from galls through a stellate distal opening in June-July and migrate
to found colonies on roots of Labiatae. Throughout Europe and in North
Africa, southwest and Central Asia, and western Siberia (Heie, 1980). 2n = 28.

Kaltenbachiella spinosa Akimoto Galls on Ulmus japonica are like those of

K. japonica (see above), but generally with larger, thicker spines arranged more
irregularly (Fig. 134J, p. 898; and Akimoto, 1985a). Emigrant alatae (BL 1.0-
1.4 mm) leave galls in July-August for an unknown secondary host. As yet only
known from Japan. 2n = 18 (Akimoto, 1985a; Blackman, 1986, as japonica).

Kaltenbachiella ulmifusa (Walsh and Riley) Galls on Ulmus rubra are large,
spindle-shaped, bag-like, about 2.5 cm long, green when young and becoming
730 Kurisakia

straw-coloured when mature, projecting upward from mid-rib of leaf

(Fig. 134I, p. 898; and Patch, 1910b). Emigrant alatae, BL 1.4-1.5 mm, with
media either unbranched or once-branched, leave galls in June-July to found
colonies on roots of Labiatae; successful transfers were made to Lycopus
virginicus (Smith, 1985). Distributed throughout the range of U. rubra in
North America.

KURISAKIA Takahashi Thelaxinae

About seven species in East Asia, described from Juglandaceae (or from
perhaps misidentified trees with similar leaves) and Fagaceae. Closely related
to Glyphina and in fact, although Kurisakia are generally paler and less
sclerotized than Glyphina, there are no good morphological distinguishing
features between these two genera. Many of the characters used to distinguish
between species and subspecies of Kurisakia are subject to environmental
variation, so that the separate identity of most species needs experimental con-
firmation; possibly there are only two or three valid species in the genus. There
is no information on sexual morphs or life cycles for any species of Kurisakia.
Accounts are available for Japan (Takahashi, 1960a) and China (Chang and
Zhong, 1979a).

Kurisakia ailanthi Takahashi Apterae broadly oval, colour in life not

recorded in original description, but probably greenish-yellow with green
markings, with legs and antennae pale; BL c. 2.7 mm. Described from Ailan-
thus altissima in Japan (Takahashi, 1960a), but almost certainly this was a
misidentification of Juglans sieboldiana, which has very similar leaves. Dis-
tinction from K. onigurumi, apart from body size and size-related characters,
is not clear. Sorin (1989) gave a detailed description, including dimorphic first
instars, of a subspecies, Kurisakia ailanthi sawagarumii, curling leaflets of
Pteryocarya rhoifolia in Japan. Life cycle unknown.

Kurisakia indica Basu Apterae oval, colour in life not recorded, antennae
and legs pale; BL 1.6-1.9 mm. Alatae have paired dark markings on most
abdominal tergites. On undersides of leaves of Engelhardtia spicata in
West Bengal, feeding without evident injury to host (A.N. Basu, 1968).
Similar aphids were collected in Sikkim (BMNH colln, leg. M.R. Ghosh and
S. Chakrabarti), but from an unlikely host, Fluggea macrocarpa (Euphor-
biaceae). Life cycle unknown.

Kurisakia onigurumii (Shinji), Plate 4e, f Apterae oval, body greenish-yellow

with green cross-bands, legs and antennae pale; BL 1.2-2.0 mm. Alatae have
black head and thorax and paired green markings on ABD TERG 1-6, often
fused to form cross-bands (Moritsu, 1983, p. 212). In folded leaflets of Juglans
 Lachnus 731

sieboldiana and Pterocarya rhoifolia in Japan. Specimens assigned to this

species were also collected from Pt. stenoptera in Hangzhou, China (BMNH
colln, leg. VFE). Life cycle unknown; according to Shinji (1924), the indi-
viduals produced in July are all alatae, and fly away. 2n = 18* (for specimens
from Pt. stenoptera in China).

Kurisakia querciphila Takahashi Apterae yellow, with a pair of longitudinal

green dorsal stripes; BL 1.2-2.0 mm. On Q. acutissima in Japan, often form-
ing large populations. Paik's (1965) record of K. onigurumii on Q. acutissima
and Castanea crenata in Korea should be referred to this taxon, which was
described (Takahashi, 1960a) as a subspecies of K. onigurumii, but is here
given full species status.

Kurisakia sinocaryae Chang Described from Carya cathayensis in Zhejiang,

China (Chang and Zhong, 1979a). Alate viviparae have more numerous secon-
dary rhinaria than other described Kurisakia species (distributed ANT III
39-51, IV 9-13, V 5-7), and these are not arranged in a row (i.e. the
sensoriation is as one would expect to find in a male). Jin (1982) studied the
population ecology of this aphid.

Kurisakia sinoplatycaryae Chang Described from Platycarya strobilacea in

Zhejiang, China (Chang and Zhong, 1979a). Apterae have a rugose tergum
and hairs on the hind tibia only about as long as the width of the tibia at mid-
length, and alatae have a CAUDA with a distinct basal constriction.

Kurisakia yunnanensis Chang Alate viviparae only are described, from

China, on Cassia siamea, which is unlikely to be the true host. Similar to, or
synonymous with, K. onigurumii.

LACHNUS Burmeister Lachninae: Lachnini

About 14 palaearctic and one nearctic species of medium to large aphids, the
alatae usually having pigmented wings, mostly associated with Fagaceae and
attended by ants. It is a taxonomically difficult genus, as indicated under
L. roboris. Accounts are available for Central Europe (Heinze, 1962), Poland
(Szelegiewicz, 1978), Hungary (Szelegiewicz, 1977), India (A.K. Ghosh,
1982b), Middle East (Bodenheimer and Swirski, 1957) and Japan (Sorin,
1980). Until about 1930 the species were often placed in Pterochlorus and the
name Lachnus was applied to aphids of the genus Cinara. Several species of
Pauesia are recorded as parasitoids in Asia.

Lachnus acutihirsutus Kumar and Burkhardt Apterae have black head and
thorax and brownish-black abdomen; BL 2.8-4.4 mm. Alatae have forewings
732 Lachnus

mostly pigmented, except for two large clear triangles on basal half and a
small clear spot between pterostigma and Rs. On twigs of Quercus incana and
Qu. dealbata in northern India (Uttar Pradesh, Himachal Pradesh). Sexuales
and life cycle unknown. 2n = 16 (J. Dutta and D.C. Gautam, pers. comm.).

Lachnus allegheniensis McCook (= montanus Wilson) Apterae rusty or

yellowish-brown, with legs brown to black; BL 3.5-4.5 mm. The rare alate
morph was described by Hottes (1954a; as montanus); the forewing veins are
heavily bordered with fuscous, which extends between the veins at the apex
of the wing. On twigs of Quercus spp. (gambelii, gunnisonii, virginiana),
collected in various parts of North America, but of rather sporadic occur-
rence. Monoecious holocyclic; apterous males and oviparae in October
(Palmer, 1952).

Lachnus crassicornis Hille Ris Lambers Colour of apterae in life not

recorded; BL 2.2-2.5 mm. On Quercus ithaburensis (BMNH colln, leg.
DHRL) in Israel. Other morphs and life cycle unknown. Records from Turkey
on Q. aegilops (Çanakçioglu, 1967, 1975) are of a different, undescribed
species. There is also a second undescribed Turkish species of Lachnus, on
Q. persica (BMNH colln, leg. G. Remaudière). (See Quercus key D.)

Lachnus fici Takahashi Apterae brownish-black; BL c. 4.5 mm. Alatae have

blackish-brown pigment at bases of wings, not extending to Culb in forewing,
and wing membrane otherwise only slightly dusky around margin. On bran-
ches of an unidentified Ficus sp. in Taiwan, and apparently not recorded since
the original description (Takahashi, 1933c). The synonymy with Nippolachnus
himalayensis proposed by Tao (1958) seems doubtful.

Lachnus pallipes (Hartig) (= exsiccator, iliciphilus, longirostris) Apterae

shining dark reddish- to blackish-brown; BL 3.5-4.Omm. Alatae have a pat-
tern of forewing pigmentation similar to L. roboris, but with a more extensive
clear area between Rs and media. On older branches and stems of Quercus
spp. and Fagus sylvatica, causing feeding damage by rupture of the cambium.
It may feed in ant shelters on roots of Fagus in summer (Heinze, 1962).
Monoecious holocyclic; oviparae and small apterous males occur (on both
Fagus and Quercus) in October. In northwest and Central Europe. 2n = 8
(Blackman, 1990; as iliciphilus). Szelegiewicz (1975) discussed the synonymy
of pallipes and longirostris, and described a new, closely-related species from
Quercus sp. in Korea, L. chosoni.

Lachnus roboris (Linnaeus) Apterae shining blackish-brown; BL 3.5-

5.0mm. Alatae have forewing membrane pigmented except for clear patches
on either side of Rs, a clear band running obliquely across base of media and
distal part of Cu1a, and a clear triangle from Cu lb to the wing-base. On twigs
and small branches of Quercus spp. and also Castanea vesca. In Europe east
to Ukraine, the Mediterranean region and Lebanon. Monoecious holocyclic;
apterous oviparae and large alate males in September-October. Michel (1942)
 Lachnus 733

studied its biology on oaks. Lachnus roboris is likely to be a complex of species

with different host plant associations and karyotypes (see also B & E, 1984,
p. 290). Chromosome numbers ranging from 7 to 17 are recorded (Blackman,
1990).

Lachnus salicis Chakrabarti and Raha Colour in life not recorded, probably
brownish; BL of aptera 3.5-5.4mm. Alatae have clear wings. In large dense
colonies on stems of Salix babylonica and S. tetrasperma, attended by ants.
Sexual morphs not recorded; apterous viviparae and immatures may over-
winter in bark crevices (Chakrabarti and Raha, 1988). In Uttar Pradesh, India.
Possibly a synonym of L. longirostrum, a very similar aphid described from
Salix fragilis in Himachal Pradesh (David and Ghosh, in A.K. Ghosh, 1982b).
It is likely that these willow-feeding aphids, with general appearance and
biology very like Tuberolachnus salignus but lacking a dorsal tubercle, are
wrongly placed in Lachnus.

Lachnus shiicola Sorin Apterae shining brownish-black; BL 3.7-4.0 mm.

Forewing pigmentation of alata similar to that of L. tropicalis. On Castanopsis
sp. (?cuspidata) in Japan, feeding in scars of the bark of the trunk near its
base, and on exposed roots of old trees, covered by ant shelters. Sexuales and
life cycle unknown. Closely related to L. tropicalis, but rather longer-haired
and with shorter antennae and HT II (Sorin, 1980).

Lachnus swirskii Hille Ris Lambers Apterae reddish-brown, broadly pear-

shaped; BL 3.1-4.4mm. Alatae have forewing pigmentation closely resem-
bling that of L. roboris. On branches of young Quercus spp. (aegilops,
ithaburensis), recorded from Israel and Turkey (Hille Ris Lambers, 1954b;
Çanakçioglu, 1967, 1975). Sexuales and life cycle unknown. Davatchi et al.
(1957) described a form with longer hairs on the vertex, collected from
Q. persica in Iran, as a subspecies, L. swirskii persicae.

Lachnus takahashii Sorin Apterae brownish-black, not shiny; BL 3.8-

4.3 mm. Forewings of alata mainly infuscated, but with several clear patches
of irregular shape. On Quercus acutissima in Japan, feeding in scars on the
bark of old trees, usually on the basal part of the trunk in ant shelters (Sorin,
1980). Sexuales and life cycle unknown.

Lachnus tatakaensis Takahashi Apterae brownish-black, with many large

black circular spots in rows on dorsal abdomen. Antennae black, legs reddish-
brown and black; BL 5.5-5.7 mm (Takahashi, 1937b). Alatae not described.
On trunk and branches of an unidentified Salix sp. at high altitude in Taiwan.
Life cycle unknown. Possibly this species belongs in either Tuberolachnus or
Pterochloroides.

Lachnus tropicalis (van der Goot) (= japonicus Matsumura) Apterae shin-

ing brownish-black; BL 3.8-5.1 mm. Alatae have forewing wholly infuscated,
except for a clear patch between pterostigma and Rs, and a clear band from
734 Lambersaphis

base of media to distal part of Cula. On twigs and stems of Quercus spp. and
Castanea spp. in East and Southeast Asia (India, Japan, Korea, China,
Siberia, Java, Malaya). Dense populations of a Lachnus that appears to be
this species were also collected on twigs of Casuarina equisetifolia at high
altitude in New Guinea (BMNH colln, leg. J. Szent-Ivany). Records from
Castanopsis (= Shiia) are possibly all L. shiicola. Monoecious holocyclic in
Japan, with apterous oviparae and alate males (Shinji, 1927; Moritsu, 1983);
possibly anholocyclic in warmer regions. Lachnus siniquercus Zhang, described
from Q. liaodongensis (?liaotungensis Koidzumi) in China (Zhang and Zhong,
1982b) is closely related and could be large specimens of tropicalis. The
tropicalis/shiicola group perhaps involves a complex of species or races,
paralleling that thought to occur in L. roboris. B.C. Das and Chakrabarti
(1989a) recorded a new parasitoid from L. tropicalis, in India. 2n = 12, 13 or
16 (Blackman, 1986).

Lachnus wichmanni Hille Ris Lambers Apterae dull blackish with greyish
waxy markings; BL 3.3-4.6 mm. Sides of head and sections of the femora and
tibiae are orange-brown. Alatae have the forewing membrane infuscated
distally, and very thick bands of fuscous along Rs, media and Cu1b. On trunk
and branches of Hippophae rhamnoides, attended by ants. Recorded from
Germany and Italy. Monoecious holocyclic; apterous oviparae and small
apterous males in August in southern Germany (Hille Ris Lambers, 1956c).

Lachnus yunglongensis Zhang Known only from apterous viviparae (BL

c. 4.6mm) collected on Salix sp. in China (Zhang and Zhong, 1985e). Biology
unknown. This species, like L. longirostrum and L. salicis described from
Salix in India, may have greater affinity with Tuberolachnus than with
Lachnus.

LAMBERSAPHIS Narzikulov Chaitophorinae

A genus for one species on Populus, differing from Chaitophorus in the very
short processus terminalis, sparse short needle-like dorsal hairs, crater-like
siphunculi without reticulate sculpturing and semicircular CAUDA. The
alatae have thickly fuscous-bordered wing veins like the North American
Chaitophorus populicola (which also has a rounded CAUDA).

Lambersaphis pruinosae (Narzikulov) Appearance in life unknown, prob-

ably brownish; BL of aptera c. 1.6 mm. On young shoots of Populus pruinosa
and P. diversifolia in Central Asia (Kazakhstan, Tadzhikistan; Aibasov, 1971;
Narzikulov, 1957, 1961b). Oviparae and alate males in October (BMNH colln,
leg. M.N. Narzikulov).
 Lithoaphis 735

LATGERINA Remaudière Drepanosiphinae: Phyllaphidini

One or two species on Alnus in Mexico, characterized by the apterae with

frontal and marginal multidigitate hair-bearing processes. See Remaudière
(1981) for all available information on the genus.

Latgerina orizabaensis Remaudière Apterae flattened dorsoventrally, pale

yellow with black antennae; BL 1.6-2.2 mm. Alatae have black head, thorax
and appendages and black dorsal abdominal markings. On undersides of
leaves, especially on young branches, of Alnus acuminata ssp. arguta in
Mexico (Remaudière, 1981). A more pigmented form with paler antennae
collected on A. firmifolia was described as a subspecies, L. orizabaensis
mexicana; the host plant was given in the original description as A. acuminata,
but was corrected in Remaudière and Muñoz Viveros (1985a, p. 78). Mono-
ecious holocyclic; oviparae and alate males produced by both subspecies in
September.

LITHOAPHIS Takahashi Hormaphidinae: Nipponaphidini

Two little-known species on Fagaceae, tentatively placed together because they

have heavily sclerotized apterae with fused head, thorax and ABD TERG 1-7,
and siphuncular pores.

Lithoaphis lithocarpi (Takahashi) Apterae black, rather densely covered

with white secretion. Body almost circular, dorsally flattened; BL c. 0.75 mm.
On undersides of leaves of Lithocarpus sp. in Taiwan (Takahashi, 1929; as
Astegopteryx lithocarpi). Other morphs and life cycle unknown.

Lithoaphis shiiae Takahashi Apterae black, shining. Body almost circular,

strongly convex dorsally; BL 1.2-1.5 mm. Alatae have secondary rhinaria
distributed ANT III 21-23, IV 10-11, V 8-11. Apterae are found on branches
of Castanopsis cuspidata in Japan; immature alatae feed along the mid-ribs
on the undersides of leaves. Life cycle unknown; apparently anholocyclic on
Castanopsis in Japan (Takahashi, 1959a).
736 Lizerius

LIZERIUS Blanchard Drepanosiphinae: Phyllaphidini

A South American genus related to Paoliella, with no clear host relationships.

Three species are known to be tree-dwelling, one on Gombretaceae and two
on Lauraceae, but several other species are only known from trapped alatae.
Quednau (1974) reviewed the genus and provided keys to all known apterae
and alatae.

Lizerius braziliensis Quednau Apterae yellow-green, with long marginal

processes; BL 1.2-2.1mm. Alatae have dark head, pterothorax, antennae
and legs. On young shoots and suckers of Terminalia australis in Brazil and
Uruguay. Apterous males and oviparae in January (Quednau, 1974; and
BMNH colln, leg. VFE).

Lizerius ocoteae Blanchard, Plate 7c, d Apterae dark olive to brownish-

black, thickly coated with white wax; BL 1.6-2.0 mm. Alatae are brownish-
black with a sparse covering of wax meal. In dense colonies on young stems
and leaves of Ocotea acutifolia in Argentina (Blanchard, 1923), and Uruguay
(BMNH colln, leg. VFE). Also recorded from Phoebe porphyria (Blanchard,
1944). Alatae have been trapped in Brazil. Blanchard (1923) found oviparae
and alate males within the colony on Ocotea, but gave no collection date.

Lizerius tuberculatus (Blanchard) Apterae brownish-black; BL 1.2-1.5 mm.

Alatae brownish-black, weakly pruinose. On young growth of Nectandra sp.
(Blanchard, 1939) in Argentina and on Ocotea ?glomerata in Venezuela
(BMNH colln, leg. M. Cermeli). Alatae have also been trapped in Brazil and
Jamaica. Sexuales and life cycle unknown.

LONGISTIGMA Wilson Lachninae: Lachnini

Three species have been described in this genus, one in North America and
the others in the Himalayas and Taiwan. Very large, bark-feeding aphids,
characterized in the alatae by the elongate pterostigma extending around tip
of forewing. The differences between the species are not clearly defined and
it seems possible that they could even represent populations of one poly-
phagous species introduced to various parts of the world.

Longistigma caryae (Harris) Apterae pale brownish-grey, slightly pruinose,

with conspicuous rows of dark dorsal spots and dark siphunculi; BL 5.1-
6.6mm. On bark of numerous tree species in North America. Bissell (1978)
 Macchiatiella 737

listed 292 host records from trees in 24 genera and 16 families. Monoecious
holocyclic, with alate males, in northern USA (Washington, DC; Wilson,
1909), anholocyclic in southern states (Bissell, 1978). Marked fluctuations in
population size occur, this species being very abundant in some years (e.g.
Tissot, 1944).

Longistigma liquidambarus (Takahashi) Apterae black or brownish-black,

lightly dusted with wax powder; BL c. 7.0mm. On Liquidambar formosana
in Taiwan and Japan (Kawada and Yamashita, 1992), feeding on the bark.
Anholocyclic; no sexual morphs known. Tsumuki et al. (1993) studied its
tolerance of low temperature.

Longistigma xizangensis Zhang Apterae of BL 6.5-7.1 mm, colour in life

unknown. Recorded from several genera of trees (Salix, Populus, Prunus,
Quercus, Machilus) in Tibet (Zhang and Zhong, 1981b).

MACCHIATIELLA del Guercio Aphidinae: Macrosiphini

Two or three palaearctic species with host alternation between Rhamnus and
Polygonum. The antennal tubercles are smooth and divergent, the apterae
have secondary rhinaria on ANT III, the siphunculi are tapering and rather
short and the CAUDA is reduced to such an extent that it is almost
indiscernible.

Macchiatiella itadori (Shinji) Host-alternating between Rhamnus (japonica,

purshiana) and Polygonum spp. in Japan, China and Korea (Miyazaki, 1971).
Apterae on Rhamnus are shining yellow-brown or yellow-green with a brown
head, an extensive black dorsal abdominal patch, banded antennae and legs
and black siphunculi (for illustration see Moritsu, 1983); BL 1.8-2.1mm.
Large colonies are formed on the leaves, young shoots and woody stems,
causing leaf curl (Hori, 1927; as Acaudus rhamni). Alatae produced on
Rh. japonica were shining green according to Hori (1927), whereas those pro-
duced on Rh. purshiana were yellow (Miyazaki, 1971). Oviparae and alate
males were collected on Rh. japonica in September-October (Hori, 1927; and
BMNH colln). 2n = 12.

Macchiatiella rhamni (Boyer de Fonscolombe) Host-alternating between

Rhamnus spp. and Polygonum spp. in Europe. Apterae on Rhamnus are
yellow-brown or yellow-green, and have antennae ringed with black, pale legs
with black tarsi, pale siphunculi and variably developed black markings on
ABD TERG 3-5, which are sometimes completely absent; BL 1.9-2.3 mm. In
southern France, Italy, Portugal, Spain, and east to Turkey (BMNH colln).
Anholocyclic populations occur on Rh. alaternus in the Mediterranean area
(Barbagallo and Stroyan, 1982). A form regarded as a subspecies, M. rhamni
738 Machilaphis

tarani, alternates between Rh. cathartica and Polygonum polymorphica in

Central Asia (Nevsky, 1929a; as Neanuraphis).

MACHILAPHIS Takahashi Drepanosiphinae: Phyllaphidini

One oriental species on Lauraceae, resembling Phyllaphis and Diphyllaphis

but differing in the form of the empodial hairs and structure of the last rostral
segment (Takahashi, 1960c).

Machilaphis machili (Takahashi) Apterae elongate oval, pale green or

yellowish with waxy secretion; BL 1.6-2.0 mm. On undersides of leaves of
Machilus spp., Neolitsea sericea and Phoebe sheareri. Recorded from Japan,
(Higuchi, 1972), China, Taiwan and India (A.K. Ghosh et al., 1971d; Assam,
Meghalaya). Biology and life cycle have not been studied; sexual morphs
unknown.

MACROSIPHUM Oestlund Aphidinae: Macrosiphini

Only a few of the 120-or-so species in this genus are tree-dwelling, including
the subgenus Neocorylobium with five species associated with Betulaceae.
Sitobion is here treated as a separate genus. For an introduction to the genus
as a whole and bibliographic information see B & E, 1984, p. 294.

Macrosiphum amelanchiericolens Patch Described from Amelanchier spicata

in Maine (Patch, 1919), but not recorded since, and the types are lost
(MacGillivray, 1968). Apparently a distinctive species, with numerous secon-
dary rhinaria on ANT III in both apterae and alatae. Appearance in life and
biology unknown.

Macrosiphum californicum (Clarke) Apterae elongate spindle-shaped, green

with long, yellowish-green, dusky-tipped siphunculi; BL 1.9-2.6 mm. Alatae
are rather pale and have only 5-10 secondary rhinaria, confined to basal half
of ANT III. On young growth of Salix spp., common and widely distributed
in North America. Monoecious holocyclic; oviparae and alate males in
October (Palmer, 1952). 2n = 10.

Macrosiphum (Neocorylobium) carpinicolens Patch Apterae green with

dark legs, antennae and distal parts of siphunculi; BL 1.8-2.6 mm. On young
shoots and leaves of Carpinus caroliniana in eastern North America.
Monoecious holocyclic; Hottes and Frison (1931) described the fundatrix,
 Macrosiphum 739

alate vivipara, ovipara and alate male. Sexuales in October. See also Mac-
Gillivray (1968).

Macrosiphum (Neocorylobium) coryli Davis Apterae with reddish-brown

head and prothorax, abdomen mainly green suffused with dark brown laterally
and posteriorly, antennae and legs mainly dark, and siphunculi black con-
trasting with the pale CAUDA (Davis, 1914); BL 1.6-2.2 mm. On shoot tips
and undersides of young leaves of Corylus americana and C. cornuta (incl.
var. California) in USA (more northerly states) and across Canada. Presumably
monoecious holocyclic, but life cycle and sexuales are apparently unknown.

Macrosiphum (Neocorylobium) corylicola Shinji Apterae dull yellow to

yellowish-green, often suffused dorsally with dark brown, antennae, legs and
siphunculi mainly black, CAUDA dusky (Miyazaki, 1971); BL 2.5-3.1 mm.
On Corylus spp. and Carpinus laxiflora in Japan and Korea (Paik, 1972).
Biology and sexuales unrecorded.

Macrosiphum euphorbiae (Thomas) Apterae spindle-shaped, variably col-

oured, usually some shade of green but sometimes pink; BL 1.7-3.6 mm.
Alatae are rather pale, with only the antennae and siphunculi noticeably
darker than in apterae. A very polyphagous species, mainly on herbaceous
plants, but also occurring occasionally on numerous tree species, especially on
young growth. A nearctic species, now distributed almost worldwide. Mainly
anholocyclic; a sexual phase on Rosa spp. occurs in North America. For
further information see B & E, 1984, p. 296-297. 2n = 10.

Macrosiphum hamiltoni Robinson Apterae pale green, with antennal joints,

tarsi and tips of siphunculi dusky to black; BL 2.2-3.1mm. Originally
described from specimens collected on Humulus lupulus in Manitoba, Canada
(Robinson, 1968), but since collected from Cornus stolonifera in Manitoba
and from C. alternifolia in New Brunswick. From the dates of collection host
alternation is unlikely and Cornus is probably the usual host.

Macrosiphum manitobensis Robinson Apterae pinkish-green; BL 2.5-

2.8mm. On Cornus stolonifera in Manitoba, Canada. Robinson (1965)
described all morphs, including oviparae and alate males in late September,
but did not rule out host alternation, as no specimens were found on Cornus
in July-August. 2n = 10. [Specimens of a Macrosiphum from C. stolonifera
in British Columbia (BMNH colln, leg. C.K. Chan) appear to be another,
undescribed species.]

Macrosiphum (Neocorylobium) pseudocoryli Patch Apterae green with

antennae, tibiae and siphunculi dusky to dark, especially at apices; BL
2.4-3.6 mm (MacGillivray, 1968). On young growth and undersides of leaves
of Ostrya virginiana and Corylus spp. in northwestern USA and Ontario,
Canada. Presumably monoecious holocyclic; oviparae in Virginia in late
September (BMNH colln, leg. RLB). 2n = 10*.
740 Matsumuraja

Macrosiphum pyrifoliae MacDougall Apterae, according to the original

description, are dusky brown, with head slightly darker; BL 2.4-3.6 mm.
Described from Pyrus occidentalis (= Sorbus sitchensis) in British Columbia,
Canada (MacDougall, 1926). MacDougall counted 14-17 secondary rhinaria
on ANT III of apterae, but apterae identified as this species from the type
locality (Merritt) in MacDougall's collection have only 3-8 secondary rhinaria.
Aphids believed to be this species were collected more recently in the Van-
couver area, from Sorbus aucuparia and S. sitchensis (BMNH colln, leg.
C.K. Chan). These had green and red colour morphs (C.K. Chan, pers.
comm.). Apparently monoecious holocyclic on Sorbus, although sexuales
have not been described. Closely similar to M. euphorbiae. 2n = 10, 11 and
12*.

Macrosiphum stanleyi Wilson Apterae broadly spindle-shaped, green; BL

3.0-4.4mm. Alatae have brown-black head and thorax. On leaves and stems
of Sambucus spp. (callicarpa, cerulea, pubescens) in western North America,
from Colorado to Alaska. Monoecious holocyclic; oviparae and alate males
in Colorado in August (Palmer, 1952), but in British Columbia oviparae are
present as early as June (BMNH coll, leg. C.K. Chan). 2n = 16*. [The generic
position of this aphid is uncertain, the karyotype indicating that it may be a
Sitobion; apterae (?fundatrices) collected in March-April in British Columbia
are much more like Sitobion than later generations, having very short antennal
hairs and cylindrical siphunculi with very little distal reticulation.]

Macrosiphum tiliae (Monell) Apterae green or pink, with dark antennae,

tibiae and siphunculi; BL 2.4-2.7 mm. Colonies occur on new growth, espe-
cially suckers, of Tilia americana (Hottes and Frison, 1931). Widely dis-
tributed in northern USA. Monoecious holocyclic; oviparae and alate males
in Wisconsin in September (Davis, 1914).

Macrosiphum (Neocorylobium) vandenboschi (Hille Ris Lambers) Apterae

pale green, with black-tipped antennae, legs and siphunculi; BL 2.1-2.2 mm.
On Corylus cornuta var. californica in California (Hille Ris Lambers, 1966d;
as Corylobium vandenboschi). Biology and sexual morphs unknown.

MATSUMURAJA Schumacher Aphidinae: Macrosiphini

About ten East and Southeast Asian species, almost all on Rubus (Miyazaki,
1971), but one species is described from Ficus.

Matsumuraja calorai Calilung Apterae brownish-yellow; BL 1.1-1.3 mm.

On Ficus ribes var. cuneata in the Philippines (Calilung, 1972). Other morphs,
biology and life cycle unknown.
 Melanaphis 741

MEGALOPHYLLAPHIS Ghosh, Ghosh and Raychaudhuri

Drepanosiphinae: Drepanosiphini

An oriental genus related to Drepanosiphum, but with an elongate caudal

knob and longer dorsal body hairs. Probably the genus is associated with
Acer, but only one species has been described, Megalophyllaphis obscura from
north Bengal, India (M.R. Ghosh et al., 1971) and its host was not identified.
However, another undescribed species apparently belonging to this genus was
collected on Acer buergianum in Hangzhou, China (BMNH coll, leg. VFE).

MELANAPHIS van der Goot Aphidinae: Aphidini

About 20 palaearctic species with short siphunculi, closely related to Rhopalo-

siphum, and with the same association with Rosaceae (Pyroidea) and
Gramineae, although only a few species have proven host alternation. The
taxonomy of those East Asian species that feed on Arundinaria and related
bamboos is in need of further revision. See also B & E, 1984, p. 302.

Melanaphis arundinariae (Takahashi) Apterae purplish brown (Liao, 1976);

BL 1.2-1.7 mm. On undersides of leaves of bamboos of the Arundinaria
group (Pleioblastus, Yushania niitakayamensis) at high altitude in Taiwan.
Life cycle unknown. Specimens from India described as M. arundinariae (e.g.
Raychaudhuri and Banerjee, 1974) have apterae with 6-segmented antennae
and several other differences from the Taiwanese species, and are more
likely to be the secondary host forms of M. pahanensis. Other long-haired
Melanaphis from northern India, but with shorter siphunculi, have been
described as a distinct species, M. meghalayensis (q.v.).

Melanaphis bambusae (Fullaway) Apterae on bamboos are dark brown to

black with waxy grey dorsal abdominal markings, legs and antennae mainly
pale; BL 0.8-1.4 mm. Host-alternating in Japan between Photinia villosa
and bamboos (Sorin, 1962); elsewhere (and also commonly in Japan) popula-
tions are anholocyclic on bamboos (Arundinaria, Bambusa, Phyllostachys).
In East and Southeast Asia, Australia, India, Mediterranean area and USA
(Louisiana, Hawaii). Yi and Su (1985) studied predation by a coccinellid in
China. 2n = 8 or 10.

Melanaphis meghalayensis Raychaudhuri and Banerjee Apterae greenish to

dark brown, often dusted with wax; BL 1.3-1.9 mm. On undersurfaces of
leaves, or on stems at ground level, of unidentified grasses and bamboos in
742 Melanocallis

Meghalaya and West Bengal, India (Raychaudhuri and Banerjee, 1974). A

subspecies, bengalensis, with fewer hairs on the CAUDA and genital plate,
was recorded from Arundinaria sp. in West Bengal. Life cycle not studied.
Agarwala and Mahapatra (1990) described oviparae of M. meghalayensis from
an unidentified bamboo, indicating that this species may be partly holocyclic
and monoecious on Gramineae. However it is not clearly differentiated
morphologically from the secondary host form of M. pahanensis.

Melanaphis pahanensis (Takahashi) Host-alternating between Pyrus spp.

and Gramineae in northern India (Raychaudhuri and Banerjee, 1974; Medda
and Chakrabarti, 1992). Apterae on Pyrus in spring are dark brown dusted
with wax; BL 1.8-2.1 mm. On undersides of leaves, causing them to curl.
Originally described (Takahashi, 1950) from Malaya, as brownish-purple
apterae secreting cottony wax, BL c. 1.25 mm, on an unidentified bamboo of
the Arundinaria group.

Melanaphis pyraria (Passerini) Apterae dark brown; BL 1.3-2.1mm. On

undersides of pear leaves, rolling the leaves transversely or diagonally to
the mid-rib, attended by ants (Andrasfalvy, 1968). Host-alternating between
Pyrus communis and Gramineae (B & E, 1984, p. 303). 2n = 8.

Melanaphis siphonella (Essig and Kuwana) Apterae dark brown, lightly

dusted with white wax; BL 1.4-1.9 mm. On Pyrus pyrifolia in Japan (Essig
and Kuwana, 1918) and P. montana in Korea (BMNH colln, leg. W.H. Paik).
Biology and life cycle unknown.

MELANOCALLIS Oestlund Drepanosiphinae: Phyllaphidini

One species near Tinocallis, but with differences in the ventral structure of
the head.

Melanocallis caryaefoliae (Davis) All viviparae alate, squat-bodied, very

dark green to black with small tufts of white wax and conspicuous paired
black dorsal tubercles, those on ABD TERG 2 being especially large; BL
0.9-1.9 mm. On both upper and lower sides of leaves of most Carya spp., but
especially C. illinoensis. Feeding causes yellow spots on leaves, reducing
photosynthesis (Wood and Tedders, 1986) and large populations can cause
defoliation. Effect on yield of pecans was studied by Wood et al. (1987), and
Dutcher (1985) evaluated chemical control measures. For a general account
see Bissell (1978). 2n = 14.
 Mesocallis 743

MELAPHIS Walsh Pemphiginae: Fordini

One nearctic species related to the East Asian genera of Fordini of subtribe
Melaphidina (Nurudea, Kaburagia, Schlechtendalia) and with a similar alter-
nation between Rhus and mosses. The host association was discussed by
Moran (1989).

Melaphis rhois (Fitch) Forming closed, sac-like red and white galls (Fig. 129)
on leaflets of Rhus spp. (glabra, typhina) in North America. Emigrant alatae
(BL 1.4-1.7 mm) emerge in late August-October and fly to mosses. Anholo-
cyclic populations also occur all year round on mosses and have been intro-
duced into Europe (Sweden, UK). Hebert et al. (1991) studied genetic variation
in local populations by enzyme electrophoresis. 2n = 26*.

Fig. 129. Gall of Melaphis rhois on Rhus typhina (after Moran, 1989).

MESOCALLIS Matsumura Drepanosiphinae: Phyllaphidini

Five East Asian species principally associated with Betulaceae, resembling

Tinocallis except for the lack of spinal tubercles; probably more closely related
to Pterocallis, but with marginal hairs on ABD TERG 1-4 not duplicated.
Accounts are available for Japan (Higuchi, 1972) and India (Chakrabarti,
1988; A.K. Ghosh and Quednau, 1990).

Mesocallis alnicola Ghosh Colour in life unrecorded, probably pale yellow.

Apterous as well as alate viviparae are described; BL of apterae 1.0-1.2 mm,
of alatae 1.2-1.5 mm. Alatae have 9-11 secondary rhinaria on ANT III,
744 Mesotrichosiphum

distributed along most of length of segment. On Alnus nepalensis in

Meghalaya, India (A.K. Ghosh, 1974b); records from Carpinus faginea in
western Himalaya (Banerjee and Chakrabarti, 1991) could be M. sawashibae.
Biology and sexuales unrecorded.

Mesocallis fagicola Matsumura Very small pale yellow alatae; BL c. 0.6 mm.
ANT III with about 6 secondary rhinaria spaced out along segment. Described
from Fagus sylvatica in Japan (Matsumura, 1919) and not recorded since.
Biology and sexuales unknown.

Mesocallis obtusirostris Ghosh Alatae pale yellow; BL 1.2-1.5 mm. ANT III
with 3-7 rhinaria, mostly on basal part. On Alnus nepalensis (with one record
from Betula alnoides) in India (Uttar Pradesh, West Bengal) and Nepal
(Chakrabarti, 1988). Sexual morphs were reported by Chakrabarti (1988,
p. 69).

Mesocallis pteleae Matsumura Alatae pale yellow, with ANT HI brown,

rest of antennal flagellum banded with brown, and mainly brown tibiae
and tarsi; BL 1.2-1.6 mm. Described originally (Matsumura, 1919) from
Ptelea trifoliata, which is unlikely to be a true host. Since recorded from
species of Alnus, Betula, Corylus, Carpinus and Ostrya. In Japan and
China.

Mesocallis sawashibae (Matsumura) Alatae and apterae pale yellow, with

darker antennal joints and tarsi; BL 0.8-1.5 mm. On Carpinus cordata and
C. japonica, in Japan, Korea, China and Siberia. Oviparae in Japan in
October (Higuchi, 1972). 2n =10 (Blackman, 1986).

MESOTRICHOSIPHUM Calilung Greenideinae: Greenideini

One species close to Eutrichosiphum, but with 4-segmented antennae in the

apterae and sparse, mostly short hairs, which are blunt in apterae and acute
in alatae. The description of a second species, on Lithocarpus bennettii in
Java, has not yet been published (Noordam, in prep.).

Mesotrichosiphum uichancoi Calilung Apterae pear-shaped, colour in life

not recorded; BL c. 1.3mm. On undersides of young leaves and branch tips
of Premna odorata in the Philippines (Calilung, 1967). Life cycle unknown.
 Metanipponaphis 745

METANIPPONAPHIS Takahashi
Hormaphidinae: Nipponaphidini

About seven East Asian species distinguished from Nipponaphis by the orna-
mentation of the tergum, consisting of rounded pustules which are rather
evenly spaced and sized over the central part of the prosoma. The fused ABD
TERG 2-7 have marginal hairs, but there are no spinal hairs on ABD TERG
7 and ABD TERG 8 has 2 (or rarely, 4) hairs. Two species in Japan have been
shown to alternate between Distylium and Castanopsis (Sorin, 1987b), the
other species that probably belong in the genus are only known from Fagaceae.
Metanipponaphis is in need of revision and no key to species on Castanopsis
is attempted; A.K. Ghosh and Raychaudhuri (1973a) provided a partial key.

Metanipponaphis assamensis Ghosh and Raychaudhuri Apterae aleyrodi-

form, oval, light brown to yellow' (darker in life?); BL 1.7-1.8 mm (A.K.
Ghosh and Raychaudhuri, 1973a). On Castanopsis tribuloides in Meghalaya,
India. Other morphs and life cycle unknown.

Metanipponaphis cuspidatae (Essig and Kuwana) Host-alternating between

Distylium racemosum and Castanopsis cuspidata. The galls on the twigs of
Distylium (Fig. 130D, p. 785) are quite large (c. 4cm diameter), yellowish-
green, globular and rather soft (Takahashi, 1962a; Sorin, 1987b). Emigrant
alatae emerging in autumn (BL 1.8-2.1 mm) give rise to dark purple, lightly
wax-dusted aleyrodiform apterae (BL 1.5-1.7 mm) on branches of C. cuspi-
data. Records from other plants (e.g. Cinnamonum, Ficus, Quercus, Litsea)
are likely to be misidentifications. Alate sexuparae apparently return to
Distylium in May, but anholocyclic populations also remain on Castanopsis
throughout the year (Takahashi, 1962a). In Japan and Taiwan.

Metanipponaphis echinata Ghosh Apterae aleyrodiform, oval, 'light to

dark brown' (prepared specimens?); BL 0.7-0.9 mm (A.K. Ghosh, 1974b).
Described from Castanopsis histrix in Meghalaya, India. The pustules on the
pleuromarginal region of the prosoma are elongate, tapering to a blunt apex
and often have a hooked profile; specimens in the BMNH collection from
C. carlesii and C. sclerophylla in Hangzhou, China (leg. VFE) are rather larger
but have similar ornamentation and may be this species. The feeding position
of the Indian aphids was not recorded; the Chinese aphids were feeding on
the leaves. Other morphs and life cycle unknown.

Metanipponaphis lithocarpicola (Takahashi) Apterae aleyrodiform, almost

circular, black, usually with a little white wax marginally; BL 1.0-1.3 mm. On
undersides of leaves of Lithocarpus sp. in Taiwan (Takahashi, 1933d). Other
morphs and life cycle unknown.
746 Metathoracaphis

Metanipponaphis rotunda Takahashi Host-alternating in Japan between

Distylium racemosum and Castanopsis spp. The galls on Distylium are formed
on leaf petioles, and sometimes on the fruit. They are pale green and rather
slender, 1.7-2.8 cm long (Sorin, 1987b). Alatae migrate to (unidentified)
Castanopsis spp., producing aleyrodiform apterae which are black, almost
circular, dorsally depressed, BL 1.0-1.3 mm, and live on undersides of leaves.
Alatae (?sexuparae) are produced on Castanopsis in April in the Tokyo region
(Takahashi, 1959a). Only known from Japan.

Metanipponaphis shiicola Takahashi Apterae are aleyrodiform, black, oval,

dorsally not much depressed; BL 1.1-1.5 mm. On Castanopsis cuspidata in
Japan, living on undersides of leaves along basal part of mid-rib (Takahashi,
1959a). Other morphs and life cycle unknown.

Metanipponaphis sylvestrii (Takahashi) Apterae aleyrodiform, probably

brown to black, almost circular; BL c. 1.0 mm. Described from leaves of
an unknown host plant in Shansi, China (Takahashi, 1935d). A. K. Ghosh
(1974b) collected similar apterae in Assam, India, on Lindera sp. Probably
the species is misplaced in Metanipponaphis. Other morphs and life cycle
unknown.

Metanipponaphis vandergooti Noordam Appearance in life unknown,

probably dark; BL of aptera 1.6-1.8 mm. On Lithocarpus sundaicus in Java.
An alata collected in August had BL c. 2.2 mm and secondary rhinaria
distributed III 15, IV 7, V 3 (Noordam, 1991). Life cycle unknown.

METATHORACAPHIS Sorin Hormaphidinae: Nipponaphidini

One species on Quercus in Japan, in which the apterae have very short
unsegmented antennae and prosoma partially fused with abdominal plate; the
alatae as well as the apterae lack siphuncular pores.

Metathoracaphis isensis Sorin Apterae aleyrodiform, oval, somewhat con-

vex dorsally, strongly sclerotized, blackish-brown dusted with greyish wax; BL
c. 1.0 mm. Alatae are blackish-brown with dark forewing veins and have
secondary rhinaria distributed ANT III 40-47, IV 15-19, V 6-9. On upper
sides of leaves of Quercus gilva in Japan. Mainly or entirely anholocyclic;
apterae were found on Quercus throughout the year, a few alatae being pro-
duced in late October (Sorin, 1987a).
 Micromyzus 747

MEXICALLIS Remaudière Drepanosiphinae: Phyllaphidini

Four or more species on undersides of leaves of Quercus in Mexico, all only

known from apterae, which have 4- or 5-segmented antennae and long thick
spiculose spinal and marginal hairs. Remaudière (1982b) provided all available
information. No life cycle information is available.

Mexicallis analiliae Remaudière Apterae white to creamy-white; BL 0.7-

1.1 mm. Spinal hair-bearing processes very variably developed; at least some
are usually long and cylindrical. On Quercus spp. especially Q. rugosa, in
Mexico. Certain characters, such as the length of the last rostral segment, vary
greatly between populations (see Remaudière, 1982b). A population on a
dwarf Quercus (?frutex) was described as a subspecies, M. analiliae pumilus
Remaudière.

Mexicallis areolatus Remaudière Apterae whitish with dark green marginal

spots on each segment; BL 0.8-1.2 mm. Placed in a separate subgenus
(Anacallis) on account of the peculiar development of the frontal and marginal
processes, and lack of spinal hairs except on ABD TERG 7 and 8. On Quercus
rugosa (and one specimen only found on Q. mexicana) in Mexico (Remaudière,
1982b).

Mexicallis calvus Remaudière Apterae whitish: BL 0.7-0.9 mm. Spinal hairs

on abdomen are of remarkably different sizes; those on ABD TERG 2, 6, 7
and 8 are very long with tuberculate bases, whereas those on ABD TERG 1,
3, 4 and 5 are minute. On Quercus sp. (?mexicana) in Mexico (Remaudière,
1982b).

Mexicallis spinifer Remaudière Apterae dark green, or pale greenish-grey

with two darker green irregular lateral bands running from pronotum to ABD
TERG 5 or 6; BL 0.7-1.0 mm. Dorsal chaetotaxy shows considerable variation
(see Remaudière, 1982b). On Quercus spp. (including crassipes, mexicana,
rugosa) in Mexico. One population on Quercus sp. was considered sufficiently
distinct to be designated as a subspecies, M. spinifer longicaudus Remaudière.

MICROMYZUS van der Goot Aphidinae: Macrosiphini

Zhang, in Zhang and Zhong, 1980a, described a species in this genus

(Micromyzus hangzhouensis) from Glochidion puberum in China. Apterae
have 1-3 rhinaria on ANT III, siphunculi longer than head width across
748 Microunguis

eyes and first tarsal segments with 3 hairs; BL c. 2.6mm. Probably the
species is incorrectly placed in Micromyzus, which are typically small fern
feeders.

MICROUNGUIS Tao Hormaphidinae: Nipponaphidini

One species related to Thoracaphis, but the aleyrodiform apterae are without
dorsal hairs except on ABD TERG 8 and have very short unsegmented anten-
nae. It was erroneously transferred to Neothoracaphis by Eastop and Hille Ris
Lambers (1976).

Microunguis depressa (Takahashi) Apterae aleyrodiform, blackish-brown,

narrowly yellowish-brown marginally, without wax; BL c. 0.9mm. On under-
sides of leaves of an unidentified Quercus sp. in Taiwan (Takahashi, 1933d;
Tao, 1966, 1969). Other morphs and life cycle unknown.

MIMEURIA Börner Pemphiginae: Pemphigini

One palaearctic species related to Paraprociphilus, but the apterae on the

secondary host have 1-segmented tarsi and a very different association with
the host plant.

Mimeuria ulmiphila (del Guercio) Forming terminal leaf nests on Acer spp.,
especially A. campestre, by inhibition of shoot growth, twisting and folding
of leaves. Fundatrices are olive green-grey, covered with white wax wool; BL
3.5-4.5 mm. They give rise to numerous dark brown alatae (BL 2.6-3.3 mm),
which fly over an extended period (June-November). Apterous exules live
mainly on roots of Ulmus; they are yellow, thickly wax-powdered, BL
1.3-2.3 mm, encased singly in brown microrrhizal cysts (Marchal, 1933;
Krzywiec, 1964). Such cysts have also been found on Rubus roots close to
Ulmus (Vernon, 1957). Sexuparae are produced in autumn and return to Acer,
but anholocycly on Ulmus roots often seems to predominate especially in
Western Europe. Krzywiec (1964) suspected that anholocyclic overwintering
of immature stages might also occur on bark of Acer. Recorded from Britain,
France, Germany, Hungary, Italy, The Netherlands, Poland, Russia and
Turkey. For a full account of the life cycle in Poland and descriptions of all
stages see Krzywiec (1962, 1964).
 Mindarus 749

MINDARUS Koch Mindarinae

Five species of conifer-feeding aphids, one nearctic and four palaearctic, in

which apterae have fused head and pronotum, 3-faceted eyes, well-developed
wax glands and a blunt triangular CAUDA. Alatae have forewings with an
elongate pterostigma, tapering to a point at the wing apex, with Rs arising at
its base. Heie (1980) reviewed the European species, the biology of which was
studied in detail by Nüsslin (1910).

Mindarus abietinus Koch, Plate 2c, d Apterae yellowish-green, covered with

wax wool, with antennae and legs distinctly darker; BL 1.7-2.0 mm. Alatae
(BL 1.5-2.7 mm) have dark dorsal abdominal cross-bands. On young shoots
of Abies spp. (especially alba, nordmanniana), feeding between the needle
bases. There are only three generations per year, alate and apterous progeny
of the fundatrix producing very small apterous sexuales. The eggs, black
covered with short white strands of wax, are laid in June-July, but do not
hatch until the following spring (see Johnson and Lyon, 1988, pp. 80-81).
Throughout Europe, in western Siberia and southwest Asia (Lebanon,
Turkey), and introduced and widespread in North America. Records from
East Asia may all apply to M. japonicus. Mindarus abietinus may seriously
damage or kill young shoots, or cause deformation and loss of needles
(Nettleton and Hain, 1982; Klein, 1983), and may also affect susceptibility to
spruce budworm (Mattson et al., 1989). Amman (1963) recorded natural
enemies in North Carolina, USA, and Bradbury and Osgood (1986) studied
chemical control. 2n = 12*.

Mindarus japonicus Takahashi Apterae not described, but probably similar

in life to M. abietinus. BL of alata is c. 2.0 mm (Takahashi, 1931b). On Abies
spp. (holophylla, nephrolepis) in East Asia (India, Nepal, Siberia, Japan,
Korea).

Mindarus keteleerifoliae Zhang Appearance in life unknown. Described

from shoots of Keteleeria evelyniana in Yunnan, China (Zhang and Zhong,
1984b). Alatae have secondary rhinaria distributed ANT III 17-22, IV 2-5.
Very similar to, and possibly synonymous with, M. japonicus.

Mindarus obliquus (Cholodkovsky) Apterae greenish, covered in white wax

wool; BL 1.2-1.9 mm. Alatae similar to M. abietinus, but smaller (BL
1.0-1.6 mm); differences between the two species were described by Carter and
Eastop (1973). On Picea spp. (engelmanni, glauca, sitchensis, but not abies),
feeding between needles on new shoots in spring, but without deforming
needles. In Europe, east to Turkey, and introduced into Canada and Alaska
(BMNH colln). Life cycle like that of M. abietinus, with small apterous
750 Mollitrichosiphum

sexuales in June-July. 2n = 12, according to Robinson and Chen (1969), but

a sample from P. glauca in British Columbia, Canada (leg. C.K. Chan) had
2n = 8 (RLB; unpublished data).

Mindarus victoria Essig Apterae are a soft jade-green, covered with thick
white wax wool; BL c. 1.9mm. Alatae green, wax-covered, with narrow dark
dorsal abdominal cross-bands; BL c. 2.5 mm. On shoot tips of Abies grandis
in British Columbia, Canada. Oviparae and very small dusky apterous males
in June. Essig (1939) described and illustrated all the morphs.

MOLLITRICHOSIPHUM Suenaga Greenideinae: Greenideini

About 20 species, mostly feeding on Fagaceae, or on Alnus. Body often elon-

gate, with long siphunculi. The genus is characterized by a row of transverse
(?stridulatory) ridges on the hind tibia. The tergum is usually ornamented with
conical spinules or nodules. Raychaudhuri (1956) reviewed the few species
then recognized (as Metatrichosiphon), Takahashi (1962b) reviewed the
Japanese species and A.K. Ghosh (1974a) keyed the species placed in sub-
genus Metatrichosiphon. Keys are also available for northeast India and
Bhutan (Raychaudhuri and Chatterjee, 1980), Nepal (Das and Raychaudhuri,
1983) and Java (Noordam, in prep.) Several species were described from
unidentified hosts, so the treatment of the genus here is incomplete.

Mollitrichosiphum alni Ghosh, Ghosh and Raychaudhuri Apterae pale,

elongate-bodied; BL 2.0-2.4 mm. SIPH long, slender, yellow to yellowish-
brown, 0.52-0.85 x BL in apterae and 0.8-0.9 x BL in alatae. On Alnus spp.
(nepalensis, nitida) in India (Assam, Uttar Pradesh, West Bengal), Nepal
(BMNH colln, leg. DHRL), Tibet (Zhang and Zhong, 1981b) and China
(Zhang and Zhong, 1985f). Apart from the length of the siphunculi, which
is very variable depending on the degree of alatiformity, this species does not
appear to differ significantly from M. alnifoliae, described from A. nepalensis
by Chakrabarti and Raychaudhuri (1978), which is treated here as a synonym.
Saha and Chakrabarti (1986) described the alate oviparae and males (as
alnifoliae) on A. nepalensis in September-October.

Mollitrichosiphum godavariense Das and Raychaudhuri Apterae pale to

dark brown, elongate, collected on undersides of young leaves of an uniden-
tified Quercus sp. in Nepal. SIPH of apterae pale, 0.56-0.59 x BL. Alate
oviparae, with SIPH 0.75-0.8 x BL, collected in October (Das and Ray-
chaudhuri, 1983).

Mollitrichosiphum lithocarpi (Takahashi) Apterae elongate, yellowish-green,

with antennae and legs pale yellowish-brown; BL c. 2.5 mm (Takahashi,
 Mollitrichosiphum 751

193la). SIPH of apterae long, yellowish-green, c. 0.6 x BL. On undersides of

young leaves of Lithocarpus spp. (glabra, uraiana) in Taiwan. Alatae and
other morphs not described.

Mollitrichosiphum luchuanum (Takahashi) Apterae pear-shaped, yellowish-

brown; BL 2.0-2.3 mm. Alatae black, with long black siphunculi 0.8-
0.9 x BL (siphunculi of apterous type specimen missing). On an unidentified
Quercus sp. in Loochoo (Takahashi, 1930). Not since recorded, biology
unknown.

Mollitrichosiphum nandu Basu Apterae elongate pear-shaped, light brownish-

green mottled with darker green, antennae pale to dusky, darker at apices
of segments, legs pale brown; BL 1.9-2.6 mm. SIPH of apterae pale to
dark brown with darker tips, 0.4-0.5 x BL; SIPH of alatae blackish,
0.6-0.75 x BL. On tender parts of shoots of Alnus nepalensis, particularly the
leaf axils, often in large numbers, attended by ants (A.N. Basu, 1964).
Described from West Bengal, India. After examining type material, two other
species, M. acutihirsutum Maity and Chakrabarti (1980), described from Uttar
Pradesh, and M. buddleiae A.K. Ghosh, Banerjee and Raychaudhuri (1971c),
are considered to be synonyms. Mollitrichosiphum buddleiae was described
from a presumably casual occurrence on Buddleia in Sikkim, but specimens
from Alnus nepalensis in Nepal (Das and Raychaudhuri, 1983) and from
Alnus sp. in China (Zhang and Zhong, 1985f), were subsequently identified
as this species. Saha and Chakrabarti (1986) described the alate oviparae and
males, collected in September-October. 2n = 16*.

Mollitrichosiphum nigriabdominalis Agarwala, Mondal and Raychaudhuri

Colour of apterae in life not recorded, abdomen with a central dark patch;
BL 2.2-2.9 mm. SIPH pale, 0.5-0.6 X BL. On Quercus sp. (identified as
Q. rubra, in which case non-native) in Sikkim, India (Agarwala et al., 1982).
Other morphs and biology unknown. Very similar to M. shinjii, apart from
the dark dorsal patch, which could be a variable character.

Mollitrichosiphum nigrofasciatum (Maki) Apterae pear-shaped, pale yellow-

green with an extensive and very distinctive brown-black mark covering the
pleural areas of the thorax and abdomen on each side and joined medially
across ABD TERG 3-4 (for illustration see Moritsu, 1983); BL 1.4-2.0 mm.
Alatae have a broad brown-black patch across ABD TERG 3-5. SIPH of both
morphs black; c. 0.35 x BL in apterae and 0.5-0.6 x BL in alatae. On
Quercus spp. (formosana, glauca, serrata, variabilis) and also recorded once
from Lithocarpus sp. (Takahashi, 1931a). In Japan, China and Taiwan.
Takahashi (1962b) distinguished a population on Q. glauca in Japan as a sub-
species, M. nigrofasciatum glaucae Takahashi. Sexuales and life cycle
unknown.

Mollitrichosiphum niitakaensis (Takahashi) Apterae elongate-bodied, yellow,

with pale antennae, legs and siphunculi; BL 1.9-2.7 mm. SIPH of apterae
752 Mollitrichosiphum

0.3-0.45 x BL; SIPH of alatae dark brown, 0.5-0.6 x BL. On undersides of

leaves of Quercus sp(p). in Taiwan (Takahashi, 1937b) and on Castanopsis
sp. in Java (as javanicum Raychaudhuri). Antennae of apterae can be either
5- or 6-segmented; apterae with 5-segmented antennae were originally
described as Eutrichosiphum elongatum (Takahashi, 1940), which is here
regarded as a synonym. Sexuales and life cycle unknown.

Mollitrichosiphum shinjii Raychaudhuri, Ghosh, Banerjee and Ghosh Col-

our in life not recorded; BL of aptera c. 3.0mm. SIPH of aptera pale with
dark apices, c. 0.6 x BL. Described from Quercus sp. in West Bengal, India
(Raychaudhuri et al., 1973). Very similar to M. nigriabdominalis and
M. godavariense, either of which could prove to be synonyms.

Mollitrichosiphum taiwanum (Takahashi) Apterae broadly pear-shaped,

yellow, antennae and legs yellow with darker apices; BL 1.7-2.0 mm. SIPH
of apterae yellow with black apices, 0.4-0.5 x BL; SIPH of alatae black,
c. 0.8 x BL. On Meliosma spp. (rigida, rhoifolia) in Taiwan (Takahashi, 1921;
Raychaudhuri, 1956). Closely related to M. yamabiwae on Meliosma in Japan,
but apparently separable by the key characters given. Sexuales and life cycle
unknown.

Mollitrichosiphum tenuicorpus (Okajima), Plate 12e, f Apterae elongate

pear-shaped, pale brown with pale legs and antennae; BL 1.9-2.5 mm. SIPH
of apterae dark brown, c. 0.8 x BL. Alata brown-black, very narrow-bodied,
with black SIPH about as long as body. On young shoots of Castanopsis spp.
and Lithocarpus spp. in India, Java, Taiwan, Thailand and Japan. Also
recorded from Castanospermum sp. in Sikkim, India (A.K. Ghosh and
Raychaudhuri, 1968a) and from Castanea crenata in the Philippines (BMNH
colln, leg. V.J. Calilung). There are also a very few records from unidentified
Quercus spp., but Quercus does not seem to be favoured as a host plant. Alate
sexuales are produced in Japan in autumn and winter (Takahashi and Sorin,
1959). Agarwala et al. (1987) described a new species of parasitoid from this
aphid.

Mollitrichosiphum yamabiwae Suenaga Apterae yellow-brown; BL c. 2.1

mm. Alatae have dark brown dorsal abdomen and large paired ventrolateral
black marks on abdominal segments 3-6 (Takahashi, 1962b). SIPH dark,
c. 0.6 x BL in apterae and 0.7 x BL in alatae. On Meliosma spp. in Japan
(Suenaga, 1934). Alate sexuales are produced in summer (Takahashi, 1962b).
See also M. taiwanum.
 Monellia 753

MONAPHIS Walker Drepanosiphinae: Phyllaphidini

A genus for one large palaearctic species with many distinctive features,
including a long tapering antennal processus terminalis, feeding on Betula. See
Stroyan (1977) for a generic diagnosis.

Monaphis antennata (Kaltenbach) All adult viviparae alate, robust, ven-

trally flattened, green, with very long thick black antennae and an elongate
dark pterostigma in the forewing; BL 3.3-4.3 mm. Living solitarily on Betula
spp. in Europe and eastward across Siberia to China and Japan. Immatures
are cryptic and usually press themselves close along the mid-ribs on the upper
sides of leaves. Monoecious holocyclic; green apterous oviparae and red alate
males occur in September-October (Ostanin, 1976). 2n = 20.

MONELLIA Oestlund Drepanosiphinae: Phyllaphidini

Four North American species on Carya, resembling Monelliopsis but folding

their more rounded wings flat over the abdomen when in repose. Bissell (1978)
comprehensively revised the genus, including much morphological data and
keys to alate viviparae and oviparae.

Monellia caryella (Fitch) All viviparae alate, pale lemon-yellow to greenish-

yellow, with banded antennae, seasonally variable; generations from midsum-
mer to autumn bear a continuous broad black band running around front and
sides of head and down sides of body as far as ABD TERG 3, as well as a
broad brown-black band along the anterior margin of the forewing. BL
0.9-2.2mm. On leaves of Carya spp., especially C. illinoensis and C. cordi-
formis. Monoecious holocyclic; sexuales in mid-October to early December
(Mansour and Harris, 1988). Widespread in USA (see Bissell, 1978), in
Ontario, Canada, and introduced into Israel where it is a serious pest of pecan
and biocontrol has been attempted (Mansour et al,, 1988). Smith and Severson
(1991) studied chemical recognition of the host plant, Alverson and English
(1990) studied population dynamics and Edelson and Estes (1987) studied
natural enemies in South Carolina. According to Bissell (1978), W.M. David-
son's (1914) account of M. caryella in California applies to a species of
Monelliopsis, perhaps bisetosa, and Richards' (1965) description and figures
under the name caryella apply to Monelliopsis nigropunctata. 2n = 18.

Monellia hispida Quednau All viviparae alate, pale yellow or colourless with
banded antennae, body often with 4-6 faint yellow spots internally, and later
754 Monelliopsis

generations with a broad dusky stripe running around front and sides of body,
as in M. caryella but much paler; BL 1.2-2.0 mm (Bissell, 1978). Monoecious
holocyclic on Carya spp. in eastern North America; sexuales in October-
November.

Monellia medina Bissell All viviparae alate, bright yellow with banded
antennae, and summer/autumn generations with a heavy black band running
around front and sides and along leading edge of forewing as in M. caryella;
BL c. 2.3 mm. Recorded from Carya spp. in Maryland and Ohio, USA
(Bissell, 1978). Very similar to M. caryella. Sexuales undescribed.

Monellia microsetosa Richards All viviparae alate, pale yellow, summer

generations developing 2-3 pairs of conspicuous spots of intense yellow inter-
nal pigment, especially visible through ABD TERG 1/2 and 5/6 (Bissell,
1978). Antennae are banded and leading edge of forewing is spotted with
black. BL 1.4-2.0 mm. On leaves of Carya spp., especially C. glabra, in
eastern North America from Quebec to Florida. Sexuales in September-
November. 2n = 18.

MONELLIOPSIS Richards Drepanosiphinae: Phyllaphidini

Five North American species associated with Juglandaceae, mostly on

Juglans; except for M. pecanis on pecan, visits to Carya spp. are possibly only
'casual'. Similar to Monellia, but holding their wings vertically at rest (as do
most aphids). Richards (1966a) reviewed the genus.

Monelliopsis bisetosa Richards All viviparae alate, yellowish with banded

antennae, dusky brownish tibiae and tarsi and a dark dorsal spot near apex
of femur (Richards, 1966a); BL 1.2-1.5 mm. Described from Juglans sp.,
'probably rupestris', in Mexico and also recorded, perhaps as a casual occur-
rence, on Pterocarya fraxinifolia (Walker et al., 1978). Biology and sexuales
unknown.

Monelliopsis caryae (Monell) All viviparae alate, yellow with banded anten-
nae and, in fully pigmented specimens, a dusky median streak on the head
and small dusky spots at bases of dorsal abdominal hairs (Richards, 1965);
BL 1.2-1.8 mm. Immatures have black patches at bases of dorsal and marginal
hairs. On leaves of Juglans nigra, widely distributed in North America.
Oviparae in Utah in October (BMNH colln, leg. G.F. Knowlton). 2n = 18.

Monelliopsis nigropunctata (Granovsky) (= pleurialis Richards) All vivi-

parae alate, yellow. Antennae ringed with black, and well-pigmented speci-
mens in later generations have dark sides to prothorax, dark spots at bases
 Monzenia 755

of dorsal abdominal hairs and a dark ventral spot near apex of hind femur
(Richards, 1965); BL 1.2-1.5 mm. Immatures have dark spots at bases of
dorsal hairs. On leaves of Juglans spp. and Carya spp., widely distributed in
North America. Most records of this species on Carya illinoensis should prob-
ably be referred to M. pecanis (see below). Sexuales apparently unrecorded.
2n = 10.

Monelliopsis pecanis Bissell All viviparae alate, yellow, with darker mark-
ings (when developed) similar to M. nigropunctata; BL 1.2-1.7 mm. On leaves
of Carya illinoensis in USA, Mexico and introduced to Egypt (BMNH colln,
leg. A.A. Attia) and South Africa. Sexuales, and differences from M. nigro-
punctata, with which it was confused for many years, were described by Bissell
(1983). Alverson and English (1990) studied population dynamics, Edelson
and Estes (1987) studied natural enemies in South Carolina, and de Villiers
and Viljoen (1987) investigated control with systemic insecticides in South
Africa.

Monelliopsis tuberculata Richards All viviparae alate, yellow, dark mark-

ings when developed as in M. nigropunctata; BL 1.0-1.2 mm. Described from
Juglans sp., 'probably rupestris', in Mexico (Richards, 1966a) and also
recorded from J. nigra in New Mexico, Idaho and Utah (BMNH colln).
Oviparae in Utah in October (BMNH colln, leg. G.F. Knowlton).

MONZENIA Takahashi Hormaphidinae: Nipponaphidini

A genus for one oriental species monoecious on Distylium.

Monzenia globuli (Monzen) Forming small yellow-green spherical galls on

twigs of Distylium racemosum, usually in leaf axils (Fig. 130C, p. 785; and
Moritsu, 1983, as Nipponaphis globuli). Monoecious holocyclic. According
to Sorin (1960), there are only four generations per year in Japan. Alate
sexuparae (BL c. 1.4 mm) emerge in late October and produce sexuales on
undersides of leaves. Eggs are laid on the twigs and do not hatch until early
September of the following year, the fundatrices mature rapidly and produce
one generation of apterae in the galls, which then produce the sexuparae (see
also Nishitani and Ito, 1991). However, there are also reports that alatae
emerge from galls twice a year, in June and October (Monzen, 1954). Ito and
Hattori (1983) reported gall predation by Nola innocua (Lepidoptera). In
Japan and Korea (Paik and Choi, 1969).
756 Mordvilkoja

MORDVILKOJA del Guercio Pemphiginae: Pemphigini

One North American species in which the alatae from galls have antennae with
processus terminalis longer than base and bearing 'clear spots' of unknown
function.

Mordvilkoja vagabunda (Walsh) Host-alternating between Populus spp. of

the deltoides group and Lysimachia spp. Large, irregular, multilobed galls are
formed from the stipules of cottonwood leaves (Fig. 131D, p. 803; and Ignoffo
and Granovsky, 1961b). Emigrant alatae (BL 1.8-2.4 mm) emerge from galls
in May-June and found colonies on stems, leaves or roots of Lysimachia
(Smith, 1971). Alate sexuparae returning to Populus in September-November
are often larger than the spring migrants (2.1-2.9 mm) and have a shorter,
more normal processus terminalis. Ignoffo and Granovsky (196la) described
the gall generations and Smith (1971) gave a full account of the life cycle.
2n = 20.

MORITZIELLA Börner Phylloxeridae

Two species on Fagaceae, distinguished from palaearctic Phylloxera by the

absence of abdominal spiracles on segments 2-5 and by the presence of
numerous well-developed, pigmented dorsal tubercles. The generic characters
of Moritziella do not, however, provide a satisfactory distinction from North
American species described in Phylloxera and some revision of generic con-
cepts in this group is necessary (see Holman, 1974, p. 43).

Moritziella castaneivora Miyazaki Apterae pear-shaped, broadest anteriorly,

yellowish-brown to purplish-brown; BL 0.9-1.3 mm. Feeding on fruits of
Castanea crenata in Japan (Miyazaki, 1968). Yanagibashi and Nakagaki
(1985) studied the influence of temperature on development of the apterae.
Other morphs and life cycle unknown.

Moritziella corticalis (Kaltenbach) Apterae pear-shaped, broadest anteriorly,

green or brownish, with dark brown dorsal tubercles; BL 0.6-0.8 mm. On
Quercus robur and Q. petraea, colonizing bark of previous year's growth
(Heinze, 1962) and sometimes causing serious dieback of shoots (Barson and
Carter, 1972). In Europe, and introduced to New Zealand (Sunde, 1974). Life
cycle unclear, but probably mainly anholocyclic; alate sexuparae may be
produced in summer and autumn, but no sexuales have been recorded
(Barson and Carter, 1972). Moritziella corticalis is barely distinct from some
 Muscaphis 757

little-known North American species described in the genus Phylloxera (e.g.

davidsoni, querceti), and may be an introduced North American species that
has lost its primary host (Carya; see Stoetzel, 1985b).

Moritziella sp. Apterae pear-shaped, dirty yellow to dark brown; BL 0.7-

1.0mm. On Quercus virginiana in Cuba, colonizing apical parts of young
shoots, mainly on the stem but also on young leaves which may be slightly
curled or rolled. Perhaps an anholocyclic race of a North American species
described in another genus (Holman, 1974).

MUSCAPHIS Börner Aphidinae: Macrosiphini

(= Toxopterella Hille Ris Lambers)

Seven or eight species host-alternating between Pyroidea and mosses, or

known only from either the primary or the secondary host. They are Myzus-
like aphids perhaps closest to Nearctaphis; fundatrices are densely hairy, and
give rise in the second generation to alatae which have irregular dorsal
abdominal markings and the forewing media either once-branched or with a
second branch close to the wing apex. The hind tibiae have a row of widely-
spaced Toxoptera-like ?stridulatory pegs. The host alternation was only
recently established, for M. mexicana (Remaudière and Muñoz Viveros,
1985b), the forms on Pyroidea being previously described in Toxopterella.
Remaudière and Muñoz Viveros (1985b) keyed the primary host morphs. Heie
(1992) reviewed the European species.

Muscaphis canadensis (Hille Ris Lambers) Apterae (fundatrices) broadly-

oval, colour in life not recorded but probably dark, with dark antennae and
legs and black siphunculi; BL 2.1-2.8 mm (Hille Ris Lambers, 1960c). Rolling
leaves of unidentified Crataegus sp(p). in eastern North America (recorded
from New Brunswick and Ontario, Canada, and North Carolina, USA). The
second generation are alatae (BL 1.6-2.0 mm) and migrate to an unknown
secondary host (probably a moss).

Muscaphis drepanosiphoides (MacGillivray and Bradley) Apterae (fun-

datrices) plump-bodied, matt dark brown to purplish-black with rather shiny
black siphunculi; BL 3.6-4.3 mm. In rather tightly curled leaflets of Sorbus
spp. (americana, aucupariae, sitchensis), usually with one fundatrix and its
progeny in each leaflet (MacGillivray and Bradley, 1961). The second genera-
tion are all alate (BL 1.8-2.7 mm) and migrate to an unknown secondary host,
probably a moss. Apparently of holarctic distribution; across Canada (British
Columbia, New Brunswick), Finland (BMNH colln, leg. O. Heikenheimo),
Norway (BMNH colln, leg. H. Tambs-Lyche), Estonia, Russia (Leningrad),
the former Czechoslovakia. Shaposhnikov (1963) described specimens on
758 Myzocallis

S. aucupariae from Leningrad and Estonia as a subspecies, drepanosiphoides

irae, but there seem to be no consistent differences between Canadian and
European populations. Oviparae likely to be those of M. drepanosiphoides
(?irae) were collected on S. aucupariae in September in the former
Czechoslovakia (see Remaudière and Muñoz Viveros, 1985b). 2n = 12*.

Muscaphis mexicana Remaudière and Muñoz Viveros Apterae (fundatrices)

plump-bodied, velvety dark brown to reddish-brown with black head and
siphunculi; BL 1.6-2.4 mm. Their immature progeny are light brown. On
Crataegus pubescens (= mexicana) in Mexico, distorting and reddening
young leaves. Heteroecious holocyclic; the second generation are alatae (BL
1.7-2.1mm) and migrate to mosses. Gynoparae and males fly back to
Crataegus in November-December; Remaudière and Muñoz Viveros (1985b)
described the life cycle, and recorded heavy predation of spring colonies on
Crataegus.

Muscaphis smithi (Hille Ris Lambers) Apterae (fundatrices) are very plump-
bodied, dark brownish-black with a reddish tinge, with jet black siphunculi;
BL 3.0-4.0 mm. In tightly curled and reddened leaves of Malus angustifolia,
attended by ants. In North Carolina, USA (Hille Ris Lambers, 1962a).
Immature progeny are reddish, all developing into reddish emigrant alatae (BL
1.7-2.1mm), which fly away in July to colonize an unknown secondary
host - probably a moss.

Muscaphis stroyani (Smith) Apterae (fundatrices) black; BL c. 2.5 mm. On

Crataegus punctata in North Carolina, USA (Smith, 1980). Dark reddish
alatae (BL 1.6-2.0 mm) are produced in the second generation, in June-July.
Life cycle unknown; probably host-alternating to mosses.

MYZOCALLIS Passerini Drepanosiphinae: Phyllaphidini

About 37 species of small, delicate, usually yellowish aphids with a knobbed

CAUDA and bilobed anal plate, mostly monoecious holocyclic on Fagaceae.
Immatures usually have long, capitate dorsal hairs. Adult viviparae are all
alate, except in a few species. Forewings are variably pigmented, but there is
generally at least a dark spot at the base of the pterostigma. Quednau and
Remaudière (1994) revised the world fauna and provided a subgeneric classifi-
cation and keys to species. The main regional accounts are by Stroyan (1977,
Britain); Heie (1982, Fennoscandia and Denmark); Higuchi (1972, Japan);
Richards (1965, Canada); Boudreaux and Tissot (1962, the subgenus Lineo-
myzocallis on North American red oaks) and Quednau and Remaudière (1987,
the subgenus Castaneomyzocallis on North American chestnuts). Hoplocallis
is here treated as a separate genus. Trioxys pallidus and Praon flavinode occur
 Myzocallis 759

in Myzocallis spp. as well as in the related genera Chromaphis, Eucallipterus,

Tinocallis and Tuberculatus.
Myzocallis (Californicallis) agrifolicola Richards Alatae have black head
and thorax, abdomen either reddish-brown or yellowish with a pattern of
paired brown or black markings, antennae and legs mainly pale, and forewings
with a pattern of broad fuscous bands which only partly follow the veins; BL
2.0-2.3 mm. Immatures yellowish with dark spots at bases of dorsal and
marginal hairs. On undersides of leaves of Quercus agrifolia and Q. wislizeni
in California, USA (Richards, 1965, 1968f; Hille Ris Lambers, 1966d). Sex-
uales undescribed, biology unstudied.
Myzocallis (Lineomyzocallis) bella (Walsh) Alatae yellow with broad black
bands down sides of thorax to wing insertions and continuing along leading
edges of wings; antennae ringed with black and legs black except for bases
of femora, trochanters and coxae. BL 2.5-3.0 mm. Immatures yellow, with
legs yellow except for black tarsi and dorsum distinctively marked with 5 pairs
of black patches forming an oval shape. On various Quercus spp. through-
out eastern North America. Monoecious holocyclic; sexuales in October-
December (Boudreaux and Tissot, 1962).

Myzocallis boerneri Stroyan Alatae pale yellowish, with head and thorax
sometimes partly dusky, dorsal abdomen with small paired spinal and
marginal specks of brown pigment (visible with a hand lens), antennae ringed
with brown-black; BL 1.3-2.2 mm. Immatures are yellow with paired, dusky,
rather indistinct spinal and marginal spots. On undersides of leaves of Quercus
spp. (cerris, ilex, infectoria, suber, variabilis) in Europe, southwest Asia
(Israel, Iran, Lebanon) and introduced on European oaks to South Africa
(I.M. Millar, pers. comm.), New Zealand (Rohitha, 1982) and California.
Monoecious holocyclic; sexuales in October-November (northern hemisphere).
Very similar to, and often confused with, M. komareki or M. schreiberi (see
Hille Ris Lambers and Stroyan, 1959; López and Nieto Nafria, 1983).
2n = 14*.

Myzocallis carpini Koch Alatae pale yellow to yellowish-white, without

dorsal abdominal markings. Antennae ringed with black, and forewing with
a black spot at base of pterostigma; BL 1.3-2.2 mm. On undersides of
leaves of Carpinus betulus, especially when used for hedging. In Europe,
Mediterranean area, Middle East and introduced into New Zealand and North
America (Oregon, British Columbia). Monoecious holocyclic; oviparae in
early November in Britain and in late March in New Zealand (BMNH colln,
leg. A.D. Lowe). 2n = 14*.

Myzocallis (Castaneomyzocallis) castaneae (Fitch) Alatae yellow with black

ANT III-VI, tibiae and tarsi. The forewing has the pterostigma with a black
spot at its base and a line of pigment following its posterior margin, and vein
Culb and base of Cu1a black-bordered. BL 2.1-2.6 mm. On undersides of
leaves of Castanea dentata and Castanea sp(p). in North America (Quednau
760 Myzocallis

and Remaudière, 1987). Monoecious holocyclic; sexuales occur in October in

California (Essig, 1917).

Myzocallis (Castaneomyzocallis) castaneoides (Baker) Alatae yellow, with

ANT III-VI and tibiae entirely black or with paler sections and forewings as
in castaneae; BL 1.5-1.9 mm. On Castanea spp. in eastern USA and Mexico.
Sexuales undescribed; Tissot's (1932a) description of sexual morphs under this
name should be applied to M. tissoti (Quednau and Remaudière, 1987).

Myzocallis (Agrioaphis) castanicola Baker Alatae yellow, distinctively

marked with a dark median stripe on head and thorax and paired black spinal
and marginal patches on dorsal abdomen; BL 1.6-2.3 mm. Antennae dark
beyond the basal half of ANT III, tibial apices and tarsi dark, forewing veins
rather dark ending in fuscous spots and siphunculi dark. Immatures yellowish
with brown-black dorsal markings. On undersides of leaves of numerous
Castanea spp. and Quercus spp. In Europe, Middle East, southern Africa,
St Helena, Australia, New Zealand, South America (Brazil, Chile) and western
North America (California, British Columbia). Monoecious holocyclic;
sexuales occur on both Castanea and Quercus in October in Britain and in May
in Australia. Cottier (1953) gave a detailed account of this aphid in New
Zealand, including descriptions of sexual morphs. Quednau and Remaudière
(1994) distinguished a subspecies, leclanti, specific to Castanea sativa in
southern Europe (France, Corsica, Greece, Turkey). 2n = 14.

Myzocallis (Pasekia) cocciferina Quednau and Barbagallo Alatae pale

yellow, with head and thorax pale brownish laterally, abdomen with rather
large, pale brown, paired dorsal patches, antennae and legs mainly pale and
siphunculi pale; BL 1.4-2.4 mm. Immatures are pale yellowish with brown
dorsal markings. On undersides of leaves of Quercus calliprinos, Q. coccifera
and Q. ilex in the Mediterranean region. Sexuales unknown; it seems to be
anholocyclic, at least in Sicily (Quednau and Barbagallo, 1991). 2n = 14*.

Myzocallis coryli (Goetze) Alatae pale yellow to yellowish-white, antennae

ringed with black and forewing with a black spot at the base of the ptero-
stigma; BL 1.3-2.2 mm. On undersides of leaves of Corylus spp. In Europe,
southwest Asia, Africa (Atlas Mountains; Heie, 1982), Japan (Shinji,
1941), Tasmania, New Zealand, western North America, and South America
(Argentina, Chile). Monoecious holocyclic; sexuales in October-November in
Western Europe and in April-June in New Zealand (Cottier, 1953). Messing
and Aliniazee (1991) studied post-diapause egg development. Myzocallis
coryli has developed resistance to carbamate insecticides in North America
(Aliniazee, 1983) and biocontrol was attempted with a host-specific race of
Trioxys pallidus (Messing and Aliniazee, 1989). Aruta and Carvillo (1989)
reported a fungal attack in Chile. 2n = 14.

Myzocallis (Neomyzocallis) discolor (Monell) Alatae yellow, with yellow-

brown head and prothorax, and variably-developed brown dorsal abdomen
 Myzocallis 761

markings. The antennae are ringed with brown, the forewing membrane
usually has a pattern of infuscation, but is sometimes almost hyaline and the
siphunculi are always dark. BL 1.5-2.0 mm. Immatures yellow, pink or
brownish with darker dorsal abdominal spots (Richards, 1968f). Living singly
on undersides of leaves of Quercus spp. (mostly on white oaks), widely dis-
tributed in North America and in Cuba (Holman, 1974). Monoecious holo-
cyclic; sexuales in September-October.

Myzocallis (Lineomyzocallis) durangoensis Remaudière and Quednau

Appearance in life not observed but probably yellow with dark pigmentation
as in M. granovskyi; BL 1.7-2.5 mm. Immatures have paired dark spinal
sclerites but pale tibiae. On Quercus durifolia and Quercus sp(p). in Mexico
(Remaudière and Quednau, 1992).

Myzocallis (Lineomyzocallis) elliotti Boudreaux and Tissot Alatae yellow,

distinctively marked with black and with black legs, as in M. bella; BL not
recorded and not measurable on mounted specimens, but said to be the
smallest of the dark-legged species of Lineomyzocallis (Boudreaux and Tissot,
1962). Immatures have paired, quadrate black patches on ABD TERG 1-5,
similar to those of bella. On Quercus schumardii and Quercus sp. in Louis-
iana, USA. Monoecious holocyclic; sexuales in November-December.

Myzocallis (Lineomyzocallis) exultans Boudreaux and Tissot Alatae yel-

lowish, with black spot behind eye, broad black bands down sides of thorax
and along leading edges of wings, antennae ringed with black, fore tibiae
black, mid- and hind tibiae dusky; BL c. 1.5mm. Immatures yellow, dusted
with a greyish wax powder. On leaves of Quercus spp. (red or pin oaks) in
eastern USA (Boudreaux and Tissot, 1962; Richards, 1968f). Monoecious
holocyclic; sexuales in October (in Pennsylvania).

Myzocallis (Lineomyzocallis) frisoni Boudreaux and Tissot Alatae yellow,

marked with black as in M. bella, except that the antennae are entirely black
beyond the basal part of ANT III; BL c. 1.9mm. Brachypterous adults are
sometimes present and may have paired dark patches on ABD TERG 3-5.
Immatures have black antennae, pale legs and dark dorsal abdominal patches.
On Quercus spp. (nuttallii, palustris) in eastern USA. Monoecious holocyclic;
sexuales in October (Boudreaux and Tissot, 1962).

Myzocallis glandulosa Hille Ris Lambers Apterae yellow, with variably

developed, paired spinal dark patches or cross-bars and smaller marginal
patches, from pronotum to ABD TERG 8; BL 0.65-1.30 mm (Hille Ris
Lambers, 1948). Alatae rather squat-bodied, with banded antennae having
2-3 secondary rhinaria near base of ANT III, dark lateral spots on the pro-
thorax, and variably pigmented spinal and marginal abdominal sclerites; BL
c. 1.3mm. (Alatae have not been properly described; Richards' (1968f)
description under this name applies to M. komareki.) On undersides of leaves
of Quercus ithaburensis in Israel. The apterae are consistently the commonest
762 Myzocallis

morph (BMNH colln). Çanakçioglu (1975) records this species from Q. cerris
and Q. aegilops in Turkey, but does not specify which morph(s) were col-
lected, and perhaps had boerneri or komareki. Life cycle requires clarification;
sexuales in late autumn and winter (Bodenheimer and Swirski, 1957), and
fundatrix-like alatae in April (BMNH colln, leg. B. Halperin). 2n = 14*.

Myzocallis (Lineomyzocallis) granovskyi Boudreaux and Tissot Alatae yel-

low with very broad black bands of dark pigment running from posterior
margins of eyes along sides of thorax and leading edges of wings. Antennae
are ringed with black, femora black on distal third, tibiae and tarsi black. BL
1.8-2.3 mm. Immatures have a grey waxy bloom, dark dorsal patches and
dark tibiae. On numerous Quercus spp., usually on upper sides of leaves
(Boudreaux and Tissot, 1962). Throughout eastern USA, in Ontario, Canada
(Richards, 1965), and also collected once in Oregon (BMNH colln, leg.
DHRL). Monoecious holocyclic; sexuales in October.

Myzocallis (Pasekia) komareki (Pasek) Alatae yellow, with head and thorax
variably marked with black (typically there are fine, black lines on the pro-
thorax, roughly in the shape of a pair of rectangles) and dorsal abdomen with
a series of large, paired quadrate spinal sclerites, which have dark margins and
paler central areas, and similar-sized but paler and more uniformly-pigmented
marginal sclerites. BL 1.1-2.2 mm. On Quercus spp. (aegilops, cerris, cocci-
fera, ithaburensis) in southeastern Europe, Mediterranean area and Middle
East (Bulgaria, the Czech and Slovak Republics, Greece, Iran, Israel, Italy,
Turkey). Monoecious holocyclic; sexuales in Iran in November (BMNH colln,
leg. S.H. Hodjat). 2n = 14*.

Myzocallis (Nippocallis) kuricola (Matsumura) Alatae rather squat-bodied,

pale green (Essig and Kuwana, 1918) or red (RLB, unpublished observations),
covered with white wax; BL 1.2-1.9 mm. Forewings are patterned with very
broad bands of fuscous that follow the veins. Brachypterae occur commonly
(Takahashi, 1923). On undersides of leaves, along the mid-ribs, of Castanea
spp. and Quercus spp. In Japan, Taiwan, China and Korea. Monoecious
holocyclic in Japan; sexuales in October-November (Takahashi, 1924a;
Shibata, 1954). 2n = 14 (Shibata's record of 2n = 12 in both male and female
cells seems to be erroneous).

Myzocallis (Lineomyzocallis) longirostris Richards Alatae yellow, antennae

ringed with black, pigmentation of body and wings variable; fully-pigmented
specimens have black sides to thorax, dark fore tibiae and dark bands along
leading edges of wings, but some specimens are rather pale (Richards, 1965).
BL 1.4-1.8 mm. Immatures bright yellow with antennae ringed with black. On
red oaks in eastern North America. In Mexico, a subspecies (tepehuaensis;
Remaudière and Quednau, 1992) occurs on Q. urbani var. parvifolia.

Myzocallis (Lineomyzocallis) longiunguis Boudreaux and Tissot Alatae

yellow, distinctively marked with black as in M. bella; BL 1.5-2.0 mm.
 Myzocallis 763

Immatures are very pale, almost translucent yellow, with antennae ringed with
black. On Quercus spp. (mostly red or pin oaks) in eastern North America
(Boudreaux and Tissot, 1962; Richards, 1968f).

Myzocallis (Pasekia) mediterranea Quednau and Remaudière Alatae pale

yellow to creamy white, with dark markings; BL 1.2-1.6 mm. On Quercus
pubescens and Q. ilex, in southern France and Spain (Quednau and
Remaudière, 1994). Specimens recorded as M. komareki from Q. pubescens
and Q. canariensis in Sicily are (Barbagallo and Stroyan, 1982) also referable
to this species. Sexuales occur along with alate viviparae from November
to January, their appearance probably depending on the timing of leaf
senescence (Barbagallo and Stroyan, 1982).

Myzocallis (Lineomyzocallis) melanocera Boudreaux and Tissot Alatae

yellow, distinctively marked with black as in M. bella, except that in more
southerly populations the antennae may be wholly black; BL 2.4-2.9 mm.
Immatures are yellow without any wax bloom and have pale legs and variable
dorsal pigmentation (Boudreaux and Tissot, 1962; Richards, 1965). On
Quercus spp., mostly of the red oak group, in eastern North America.
Monoecious holocyclic; sexuales in October-December.

Myzocallis (Lineomyzocallis) meridionalis Granovsky Both apterous and

alate morphs occur. Alatae are yellow marked with black as in M. bella;
BL 1.7-1.9 mm. Apterae are yellow with black antennae, dark tibiae and black
sclerites on ABD TERG 3-5 coalesced spinally to form a 6-lobed central patch;
BL 1.7-2.1mm. On undersides of leaves of Quercus spp. (especially nigra,
schumardii) in southeast USA. Life cycle is unclear; oviparae occur, but
parthenogenetic populations persist through winter on trees which keep their
leaves in the southernmost states (Boudreaux and Tissot, 1962).

Myzocallis (Neomyzocallis) mimica Richards Alatae yellow, with antennae

ringed with black, legs pale except tarsi, dorsum with dark markings variably
developed and wings sometimes almost hyaline but usually with a distinctive
pattern of fuscous markings between veins; BL c. 1.8 mm. Only known from
Quercus macrocarpa in Ontario and Quebec, Canada (Richards, 1965, 1968f)-
Sexuales and life cycle unknown. Very similar to, and possibly synonymous
with, M. punctata and/or M. rostropunctata.

Myzocallis (Lineomyzocallis) multisetis Boudreaux and Tissot Alatae pale

yellow, distinctively marked with black as in M. bella, except that the mid-
and hind tibiae are pale with black apices; BL 1.9-2.3 mm. Immatures very
pale yellow, with black pigment usually only at apices of antennal segments.
On Quercus spp. (mostly of red oak group, but not on Q. nigra), widely
distributed in eastern North America. Monoecious holocyclic, but populations
may persist into winter on trees retaining leaves in southern states (Boudreaux
and Tissot, 1962).
764 Myzocallis

Myzocallis (Agrioaphis) myricae (Kaltenbach) Alatae yellow or orange, with

black longitudinal markings on head and thorax and paired black spinal and
marginal sclerites on ABD TERG 1-7; BL 1.8-2.0 mm. Brachypterous
specimens are common. On undersides of leaves and young growth of Myrica
gale in northern and northwest Europe (Heie, 1982). Monoecious holocyclic;
sexuales in October. 2n = 14 (Gut, 1976).

Myzocallis (Castaneomyzocallis) nanae (Tissot) Alatae yellow or greenish-

yellow with dark or banded antennae and tibiae, light brown pterothorax,
forewings as in castaneae and paired dusky spinal and marginal abdominal
patches; BL 1.4-2.0 mm. On undersides of leaves of Castanea nana in Florida,
USA. Sexuales and life cycle unknown.

Myzocallis occidentalis Remaudière and Nieto Nafria Alatae pale lemon-

yellow with dark sides to prothorax and small dusky spinal and rather larger
marginal sclerites on ABD TERG 1-7; BL 1.5-2.2 mm. On Quercus pyrenaica,
in southwest France and Spain (Remaudiere and Nieto Nafria, 1974) and on
Q. macrolepis in Italy (S. Barbagallo, pers. comm.); there is also one specimen
in the BNMH collection from Q. pubescens (leg. G. Remaudière). Monoecious
holocyclic; males in October, but oviparae unknown.

Myzocallis (Lineomyzocallis) occulta Richards Alatae yellow, antennae

ringed with black, broad black bands from posterior margins of eyes down
sides of thorax and along leading edges of wings and black fore tibiae; BL
c. 2mm (Richards, 1968f) On Quercus rubra ( = borealis) in northeast USA
(New York) and Canada (Ontario, and one record from a planted red oak in
British Columbia). Life cycle and sexuales not known.

Myzocallis (Lineomyzocallis) pepperi Boudreaux and Tissot Alatae yellow

with banded antennae and the distinctive black markings of the subgenus; BL
1.5-1.6 mm. Immatures are pale yellow without any dorsal abdominal pigmen-
tation. On Quercus laurifolia in Florida, USA. Populations on Q. castanea
and Q. crassipes in Mexico were described as a subspecies (M. pepperi turbide;
Remaudiere and Quednau, 1992). Sexuales and life cycle unknown.

Myzocallis (Pasekia) persica Quednau and Remaudière Colour of alata

in life unknown, probably rather pale; BL 1.6-2.3 mm. On Quercus persica
in Iran and Turkey (Quednau and Remaudière, 1994). Sexuales in October.

Myzocallis (Neodryomyzus) polychaeta David Colour of apterae in life

unrecorded, probably basically yellowish-brown; antennae banded, legs pale,
dorsal body with dark markings which when fully developed are quite charac-
teristic, comprising paired spinal and marginal quadrate patches with the
numerous pleural hairs in between borne on separate small rounded scleroites.
BL 1.3-1.8 mm. The alate vivipara was descibed by A.K. Ghosh and Quednau
(1990). On Quercus spp. (dealbata, semicarpifolia) in India (Himachal
Pradesh, Uttar Pradesh) and Nepal (BMNH colln). Monoecious holocyclic;
 Myzocallis 765

oviparae and males in October (Chakrabarti and Raychaudhuri, 1974).

2n = 12 (according to Khuda-Bukhsh and Pal, 1983b).

Myzocallis (Neomyzocallis) punctata (Monell) Alatae yellow, antennae

ringed with black, legs mainly pale, dark dorsal body markings often present
but very variable. Wings variably pigmented, sometimes almost hyaline in
early spring populations but later with a distinctive pattern of patchy infusca-
tion. BL 2.0-2.5 mm. Immatures have dark spinal and marginal sclerites. On
numerous Quercus spp., especially of the white oak group in eastern North
America - where it is sometimes very common in trap catches (Schultz et al.,
1985) - and introduced into western states. In California it has been collected
on many non-native oaks and also on Lithocarpus densiflora (BMNH colln,
leg. H.G. Walker). Moneocious holocyclic; sexuales occur in September-
October in northeast USA, but in California viviparae are present through the
winter. 2n = 14.

Myzocallis (Neomyzocallis) rostropunctata Richards Alatae yellow, anten-

nae ringed with black, legs mainly pale, dorsal abdomen usually without dark
markings. Forewings with black spots at base of pterostigma and at ends of
veins. BL 1.9-2.5 mm. Only known from Quercus macrocarpa in Ontario,
Canada (Richards, 1965). Very similar to the pale, spring form of M. punc-
tata; the distinction between this species, M. mimica and M. punctata needs
further study.

Myzocallis schreiberi Hille Ris Lambers and Stroyan Alatae pale yellow, with
head and thorax slightly darker; the prothorax often has short lateral streaks
of dark pigment, the dorsal abdomen has small transversely oval, often dusky
to dark, spinal spots and paler marginal sclerites, the legs are mainly pale and
the siphunculi usually dark. BL 1.3-2.2 mm. Immatures whitish to pale straw
yellow with variable pigmentation of dorsal sclerites (Hille Ris Lambers and
Stroyan, 1959). On undersides of leaves of Quercus ilex in Europe (England,
France, Italy, Greece, Yugoslavia); rarely on other oaks. Very similar to
M. boerneri; the distinction between these two species was discussed by
Barbagallo and Stroyan (1982). Sexual morphs unknown; apparently anholo-
cyclic, at least in England and Sicily. C.E.J. Kennedy (1986b) investigated
the method of attachment of the tarsi to the smooth leaves of Q. ilex.

Myzocallis (Lineomyzocallis) spinosa Boudreaux and Tissot Alatae yellow,

with antennae ringed with black, broad black bands running from posterior
margins of eyes down sides of thorax and along leading edges of wings,
all tibiae black; BL 1.6-2.0 mm. Immatures are pale yellow with paired
quadrate patches on ABD TERG 1-4 or 1-5. On leaves of Quercus spp. of
the red oak group in eastern North America. Monoecious holocyclic; sexuales
in September-October.

Myzocallis (Pasekia) taurica Quednau and Remaudière Colour of alata in

life unknown, probably pale with dark dorsal markings: BL 1.2-1.8 mm. On
766 Myzus

Quercus spp. (?calliprinos, ?robur) in Turkey (Quednau and Remaudière,

1994). Sexuales in October-November.

Myzocallis (Paramyzocallis) tenochca Remaudière and Quednau Colour of

alatae in life not recorded, probably yellowish with rather pale legs. The
forewings have a dark band along the costa, the dorsal abdomen has variably
developed spinal and marginal sclerites and the siphunculi are dark. BL
2.0-2.5 mm. Immatures have 4 pairs of dark dorsal abdominal sclerites
(Remaudière and Quednau, 1985). On Quercus crassipes in Mexico. Life cycle
and sexuales unknown.

Myzocallis (Castaneomyzocallis) tissoti Quednau and Remaudière Colour

of alatae in life not recorded, probably yellowish, with dusky, dark-banded
antennae and tibiae; forewings as in M. castaneae, but with a rather truncate
pterostigma only about five times longer than broad; BL 1.7-2.3 mm. On
Castanea nana in Florida, USA. Sexuales in November (Tissot, 1932a; as
M. castaneoides).

Myzocallis (Lineomyzocallis) walshii (Monell) Alatae bright yellow, anten-

nae ringed with black, broad bands of black pigment running down sides of
thorax and along leading edges of wings, black fore tibiae; BL 1.6-2.0 mm.
Immatures are very pale yellow with variably developed dark dorsal sclerites
(3 pairs per segment). On Quercus spp. of the red oak group, common
throughout eastern North America (Boudreaux and Tissot, 1962) and
introduced into western USA; in California it has been collected on the native
live oak, Q. agrifolia. Recently it has been found on Q. rubra planted in
France (Remaudière, 1989).

MYZUS Passerini Aphidinae: Macrosiphini

About 55 species with gibbous antennal tubercles, probably all palaearctic,

and mostly Asian, in origin. Alatae typically have a dark central dorsal
abdominal patch. Primary hosts of heteroecious species are Rosaceae, usually
Prunus. Biology and secondary host associations are diverse. Some very
polyphagous species (ornatus, persicae, antirrhinii) include trees among their
numerous host plants. Accounts are available for Central Europe (Heinze,
1961a), Japan (Miyazaki, 1971), China (Tao, 1963) and India (R.C. Basu and
Raychaudhuri, 1976a; Raychaudhuri et al., 1980b). Moritsu and Tokumoto
(1972) keyed fundatrices on cherry trees in Japan. See also B & E, 1984,
pp.310-316.

Myzus amygdalinus (Nevsky) Apterae pale green, covered with delicate

pruinose secretion, with pale, tapering siphunculi; BL 1.5-1.9 mm. Alatae
 Myzus 767

have brown head, black pterothorax and pale green abdomen with marginal
sclerites (Nevsky, 1929a). In colonies on undersides of leaves of Prunus
amygdalus and P. mahaleb in spring (April-July), causing leaves to curl
spirally, but not change colour (Vereshchagin and Narzikulov, 1961; Nar-
zikulov, 1965b). In Central Asia (Turkmenia, Tadzhikistan, Uzbekistan). Life
cycle apparently unknown.

Myzus antirrhinii (Macchiati) Apterae mid-grey-green to dark green, occa-

sionally dark red; BL 1.4-2.2 mm. On leaves and young growth of numerous
plants, including some trees, e.g. Pittosporaceae (Hymenosporum, Pitto-
sporum), in Europe and western North America (Blackman and Paterson,
1986; Blackman, 1988). Completely anholocyclic, and only producing alatae
rather sporadically. 2n = 13 or 14.

Myzus asiaticus Szelegiewicz Apterae yellow, with brownish antennae and

siphunculi; BL 1.7-2.2 mm. Alatae have shiny dark brown head and thorax
and yellow abdomen with dark brown dorsal patch. Rolling and yellowing
the leaves of Padus asiatica Kom. (= Prunus padus var. pubescens Regel and
Tiling) in Mongolia in August (Szelegiewicz, 1969). Life cycle not known.

Myzus beibienkoi Narzikulov Apterae yellowish-green with a light dusting

of wax, antennae brownish, legs and siphunculi dark-tipped; BL 1.7-2.1 mm.
The CAUDA is triangular, only about as long as R IV+V. Alatae have an
extensive dark dorsal abdominal patch on ABD TERG 3-6, often broken into
separate transverse bars on ABD TERG 5-6. On leaves and shoot tips of
Fraxinus potamophila in spring, causing slight twisting of leaves. Apparently
heteroecious holocyclic, migrating in July to an unknown secondary host.
Return migrants and oviparae were found on Fraxinus in October-November
(Narzikulov, 1957). In Central Asia (Afghanistan, Tadzhikistan).

Myzus cerasi (Fabricius) Apterae shiny, very dark brown to black, with
antennae and legs bicoloured yellow and black and siphunculi and CAUDA
wholly black; BL 1.5-2.6 mm. Alatae have a yellow-brown abdomen with an
extensive dorsal patch. In dense, often ant-attended colonies at shoot tips of
Prunus spp., especially P. cerasus and P. avium, in spring, curling leaves and
often doing severe damage. Heteroecious holocyclic, migrating to secondary
hosts in Rubiaceae (Galium), Scrophulariaceae (Veronica and, especially in
North America, Cruciferae. However, part of the population may persist
through summer on Prunus (Gilmore, 1960). In Europe, across Asia to
Pakistan, India, Siberia and Korea. Introduced into Australia, New Zealand
and North America. Rakauskas (1984) studied phenology and natural enemies
of M. cerasi in Lithuania. Apparently a complex of species or subspecies with
different life cycles and/or secondary host associations is involved (Dahl,
1968; Gruppe, 1988a); electrophoretic differences have been found between
populations on P. avium and P. cerasus (Gruppe, 1988b). North American
emigrant alatae from P. pennsylvanica and P. virginiana have a relatively
short antennal PT. In Japan, M. cerasi umefoliae occurs on P. mume and is
768 Myzus

morphologically like European M. cerasi, but does not colonize cherries and
migrates to Artemisia capillaris (Takahashi, 1965c); Myzus prunisuctus,
described from Prunus sp. in China (Zhang and Zhong, 1980a), could perhaps
be this aphid. See also B & E, 1984, p. 311. 2n = 10 (European and North
American populations) or 2n = 12 (India, possibly umefoliae; Khuda-Bukhsh
and Pal, 1986a; Kurl and Chauhan, 1988).

Myzus cornutus Medda and Chakrabarti Apterae pale, colour in life not
recorded; BL c. 2.0mm. Alatae have a dark central dorsal abdominal patch
on ABD TERG 4-6 and separate cross-bands on other tergites. Spring popula-
tions occur on Prunus cornuta in India (Uttar Pradesh), feeding on undersides
of leaves, which are rolled longitudinally and turn red or yellow (Medda and
Chakrabarti, 1986b). Apparently heteroecious holocyclic; migration occurs
during June to an unknown secondary host (Medda et al., 1986). B.C. Das
and Chakrabarti (1990) described a new parasitoid species (Aphidius staryi)
from this aphid.

Myzus ligustri (Mosley) Apterae shiny yellow, with distal parts of siphunculi
brownish-black; BL 1.0-1.5 mm. Sporadically common on privet hedges
(Ligustrum ovalifolium, L. vulgare); leaves are rolled longitudinally into nar-
row tubes and spotted with yellow. It was described from England (1841)
before the introduction of the East Asian L. ovalifolium to Europe, so pre-
sumably the native European L. vulgäre is the type host. Monoecious holo-
cyclic, with oviparae and alate males in November. In Europe and introduced
into North America. 2n = 12.

Myzus lythri (Schrank) Apterae on Prunus green to yellowish-green with

darker green longitudinal stripes; antennae, legs, siphunculi and CAUDA
pale; BL 1.8-2.0 mm. Alatae are much darker and have a large quadrate dorsal
abdominal black patch. On Prunus mahaleb in spring, in curled apical leaves.
Heteroecious holocyclic, migrating to aerial parts of Lythrum spp. and occa-
sionally to other plants in aquatic situations. There are also several records
from Epilobium spp. and one from Rhamnus pushiana (in Oregon, USA;
BMNH colln, leg. A. Gutierrez). Throughout Europe, in southwest Asia (Iran,
Lebanon) and introduced into North America. 2n = 12.

Myzus mumecola (Matsumura) Apterae very pale yellow-green to straw-

coloured with pale antennae, legs, siphunculi and CAUDA; BL 2.0-2.4 mm.
Alatae are much darker and have a broad dorsal abdominal patch with
lateral extensions on ABD TERG 3 and 4. Spring populations curl leaves of
Prunus mume and P. armeniaca (var. ansu) in Japan; also recorded from
P. armeniaca in China (BMNH colln, leg. VFE) and from Prunus sp. in Uttar
Pradesh, India (Medda et al., 1986). Life cycle and sexual morphs are
unknown; some apterae collected on Lonicera quinquelocularis in Himachal
Pradesh, India could perhaps be the secondary host form of this species
(BMNH colln, leg. A.N. Chaudhuri), but this requires experimental confirma-
tion. 2n = 12 (Chen and Zhang, 1985b).
 Myzus 769

Myzus mushaensis Takahashi Apterae bright yellow to yellowish-green with

pale antennae, legs and CAUDA; siphunculi sometimes pale brown but
usually either distally or wholly black. BL 1.7-1.9 mm. Alatae have variable
dorsal abdominal markings, often divided mesially and/or intersegmentally
and not forming a solid patch. On Prunus spp. (cherries) in Japan, Taiwan
and Korea. Feeding by fundatrices and their progeny causes the leaf lamina
to roll longitudinally upwards on both sides of the mid-rib (Moritsu and
Tokumoto, 1972; Moritsu, 1983, p. 109). Heteroecious holocyclic; secondary
host is Plectranthus japonicus (S. Hamasaki, pers. comm.). Sexuales
undescribed. Yano et al. (1984) studied predation by Elateridae.

Myzus ornatus Laing Apterae somewhat dorsoventrally flattened, pale yel-

low or green, marked dorsally with a pattern of dark green or brownish dots
and transverse flecks; BL 1.0-1.7 mm. Very polyphagous, mostly on herbs but
sometimes on trees, e.g. Catalpa, Prunus. On upper or undersides of leaves,
often feeding away from the main veins. Anholocyclic populations occur
throughout the world. B & E, 1984, pp. 313-314. 2n = 12.

Myzus padellus Hille Ris Lambers and Rogerson Apterae bright yellow
with blackish-brown head; antennae, femora and tibiae bicoloured yellow and
blackish-brown, siphunculi black. BL 2.0-2.3 mm. Alatae have a yellow
abdomen with a black dorsal patch. Forming red and yellow blistered leaf-
galls on Prunus padus in spring (Hille Ris Lambers and Rogerson, 1946). In
Europe (England, Norway, Sweden, Finland, Austria). Heteroecious holo-
cyclic, apparently migrating to Labiatae (Galeopsis spp. - records from
Norway and Sweden; Tambs-Lyche, 1957) and Scrophulariaceae (Pedicularis
verticillata, Rhinanthus sp. - specimens from Austria; BMNH colln, leg.
H.L.G. Stroyan and R.N.B. Prior). The host plant transfers to confirm the
life cycle have not been carried out, however, and sexuales are unknown.

Myzus persicae (Sulzer) Apterae varying from whitish or pale yellowish-

green to mid-green, rose-pink or red; often darker in cold conditions. BL
1.2-2.3 mm . Alatae have a dark dorsal abdominal patch. Spring colonies curl
young leaves of the primary host, Prunus persica or, in northeastern North
America, P. nigra (Shands et al., 1969). Heteroecious holocyclic, migrating to
secondary hosts in numerous plant families (Mackauer and Way, 1976; B &
E, 1984, pp. 314-315), but also anholocyclic throughout the world, mainly on
herbs but also on many tree species. Secondary colonization of young growth
of various Prunus spp. commonly occurs. In tobacco-growing areas, spring
populations on peach may include or comprise the sibling species Myzus nico-
tianae (Blackman and Spence, 1992). 2n = 12.

Myzus varians Davidson Apterae pale green or yellow-green with banded

antennae and distal halves of siphunculi contrastingly black; BL 1.7-2.3 mm.
Alatae are very dark, with a large dorsal abdominal black patch. Spring col-
onies tightly curl young leaves of Prunus persica. Heteroecious holocyclic,
migrating to Clematis spp., but populations persist on peach throughout the
770 Nearctaphis

growing season (Alma and Arzone, 1983). Presumably native to East Asia,
where it is recorded from China, Taiwan, Korea, Thailand and Japan.
Introduced into Europe and southwest Asia, where it has become a serious
pest on peaches (Arzone and Alma, 1984; Tkachuk, 1988). It is a vector of
plum pox virus (Sharka disease). Myzus varians was described from North
America, but presumably only occurs there as anholocyclic populations on
secondary hosts. (See also B & E, 1984, p. 315.) 2n = 12.

Myzus yamatonis Miyazaki Apterae black in life, with antennae and legs
pale except at apices and siphunculi and CAUDA black; BL 1.4-1.6 mm
(Miyazaki, 1971). Alatae black in life, but with variably developed abdominal
sclerotization, the central patch being irregular and with large perforations.
On Prunus spp. (cherry trees), especially P. yedoensis, spring populations
causing heavy leaf curl. Heteroecious holocyclic, migrating to Labiatae (Salvia
nipponica, Plectranthus spp.). although populations on Prunus may persist
throughout the growing season. Yano et al. (1984) studied predation by
Elateridae.

NEARCTAPHIS Shaposhnikov Aphidinae: Macrosiphini

About 12 North American species having short siphunculi ornamented with

closely-spaced rows of small spinules or nodules and a short, usually trian-
gular, CAUDA. They mostly alternate between Pomoidea as primary hosts
and Leguminosae or Scrophulariaceae as secondary hosts. Robinson (1984)
keyed and briefly reviewed the genus. Few species are well known; it is quite
likely that those Nearctaphis species recorded from one species of primary host
may also utilize other species or genera of Pomoidea. Host alternation has
only been confirmed by experimental transfers for two species.

Nearctaphis bakeri (Cowen) Spring colonies infest tips of twigs, young

leaves and blossom buds of woody Pomoidea in spring in North America
(Crataegus, Cydonia, Malus, Pyrus). Apterae are pale green to yellow-green,
sometimes pinkish anteriorly; BL 1.4-2.3 mm. Alatae have a black dorsal
abdominal patch. Heteroecious holocyclic, producing abundant alatae in the
third and subsequent generations which migrate to Leguminosae; it is an
important pest on red clover (B & E, 1984, p. 318). R.H. Smith (1923) gave
a detailed account of this species. Widely distributed in North America, and
introduced into South America, Europe, Iran, India and Japan; introduced
populations all seem to be anholocyclic on secondary hosts. 2n = 12.

Nearctaphis californica Hille Ris Lambers Described only from the secon-
dary host, Melilotus albus, in California, USA (Hille Ris Lambers, 1970c),
but apterae collected in British Columbia on Sorbus aucuparia appear to be
 Nearctaphis 771

this species (Robinson, 1984). Appearance in life unknown, adult apterae

probably rather dark with broad dark dorsal abdominal cross-bands; BL
1.5-2.0 mm (specimens from Sorbus, BMNH colln, leg. C.-K. Chan). 2n = 12*.
Hille Ris Lambers (1970c) described similar specimens from Melilotus albus in
Utah, USA, but with a solid black dorsal patch and shorter tarsi, as a sub-
species N. californica nigrescens. He assigned alatae from Crataegus sp.
in Utah, also with a solid black dorsal patch, to nigrescens, which should
probably have full species rank.

Nearctaphis clydesmithi Hille Ris Lambers Apterae and alatae on Crataegus

in spring and early summer are dark brown to black, somewhat shiny; BL
1.6-2.1mm. Apparently heteroecious, but secondary host generations are
unknown. Oviparae and alate males were collected on C. crus-galli in October
(Hille Ris Lambers, 1970c). In eastern USA (North Carolina, Maryland,
Virginia, Wisconsin).

Nearctaphis crataegifoliae (Fitch) Apterae in spring colonies on Crataegus

are greenish or reddish, BL 1.6-2.5 mm, twisting and curling the leaves and
turning them dark purple. Heteroecious holocyclic, migrating in May-June
to stem bases, runners and roots of Leguminosae (Trifolium, Melilotus,
Lathyrus). Patch (1915b) described the life cycle (as Aphis brevis). In central
and eastern North America. Aphids with a similar biology in western USA
(California, Colorado, Utah) have much darker pigmentation of head,
posterior abdomen and siphunculi, and were described (Hille Ris Lambers,
1970c) as a subspecies, N. crataegifoliae occidentalis.

Nearctaphis sclerosa (Richards) Apterae in spring colonies dark brown;

BL about 1.9mm. Apparently heteroecious holocyclic between Crataegus
douglasii and Lathyrus nutalli in British Columbia, but with an early return
to the primary host; gynoparae and oviparae were collected on C. douglasii
in early August (Richards, 1968c, as Roepkea sclerosa). Also recorded from
Colorado (Smith and Parron, 1978).

Nearctaphis sensoriata (Gillette and Bragg) Apterae in life brown to blackish;

BL 2.0-2.2 mm. On leaves and twigs of Amelanchier sp(p). in western North
America. Alatae black, with a large solid dorsal abdominal sclerite. Probably
heteroecious holocyclic with a leguminous secondary host; an aptera was
collected on Medicago hispida and autumn populations on Amelanchier com-
prise alate gynoparae, oviparae and alate males (Hille Ris Lambers, 1970c).
However, populations persist on Amelanchier into summer and host alterna-
tion needs to be confirmed.

Nearctaphis yohoensis Bradley Apterae in life are dark brown, BL 1.7-

2.2 mm, in dense ant-attended colonies on new growth and flowers of Sorbus
scopulina in British Columbia (Bradley, 1965b). Also recorded from Montana,
Manitoba, Idaho and Alaska (the latter collection from Cotoneaster acuti-
folia; Robinson, 1984). Probably heteroecious holocyclic; Robinson collected
772 Neobetulaphis

apterae of what appears to be this species from Pedicularis (Scrophulariaceae)

in Alaska, but the life cycle needs confirmation.

NEOACYRTHOSIPHON Tao Aphidinae: Macrosiphini

About seven species related to Ericolophium and Chaetomyzus, and mostly

associated with Ericaceae, but one species is described from Salix (see also
B &E, 1984, p. 319).

Neoacyrthosiphon dubium Chakrabarti and Maity Apterae pale green; BL

2.3-2.8 mm. On undersides of leaves and young growth of Salix denticulata
and Salix sp. in Uttar Pradesh, India (Chakrabarti and Maity, 1984). Monoe-
cious holocyclic, with oviparae and alate males occurring in October.

NEOBETULAPHIS Basu Drepanosiphinae: Phyllaphidini

An oriental genus on Betulaceae, nominally with four species, but the dif-
ferences between N. alba in Japan and N. pusilla in India are those to be
expected between small and large specimens respectively of the same species,
and the other two species are also doubtfully distinct from each other.
Chakrabarti (1988) and A.K. Ghosh and Quednau (1990) reviewed the three
Indian species.

Neobetulaphis alba Higuchi Apterae are pale yellow in life, BL c. 1.5 mm.
Alatae are undescribed. On Betula platyphylla var. japonica in Japan (Higuchi,
1972). Life cycle unknown. Perhaps a synonym of N. pusilla.

Neobetulaphis chaetosiphon Quednau and Chakrabarti Colour in life not

recorded, apterae presumably rather dark, brown to black with black anten-
nae, tibiae and siphunculi; BL 2.1-2.3 mm. Alatae have no dorsal black
abdominal patch. On Alnus nepalensis and Betula spp. (alnoides, utilis) in
India (Meghalaya, Sikkim, West Bengal). Life cycle is unstudied, but viviparae
have been found through the winter. As noted by Chakrabarti (1988), the
distinction betweeen this species and N. immaculata when size and seasonal
variation are taken into account needs to be verified.

Neobetulaphis immaculata Ghosh Apterae pale yellow; BL 1.2-1.5 mm.

Alatae are undescribed. Originally described from Rosaceae in Meghalaya,
India (A.K. Ghosh, 1976), but Chakrabarti (1988) recorded it from a more
likely host, Betula alnoides, in Uttar Pradesh. Life cycle unknown.
 Neocalaphis 773

Neobetulaphis pusilla Basu Apterae yellowish-green with dark green cross-

bands on thorax and abdomen; BL 1.7-2.3 mm. Alatae have a black patch
on ABD TERG 4-6 and dark siphunculi. On undersides of leaves of Betula
spp. (alnoides, utilis) and also on Alnus nepalensis, in India (Himachal
Pradesh, Sikkim, Uttar Pradesh, West Bengal). Chakrabarti (1988) described
the apterous male and ovipara from Uttar Pradesh in October, but vivipar-
ous morphs occur through the winter months in Sikkim and West Bengal.
Neobetulaphis alba in Japan may be a synonym.

NEOCALAPHIS Shinji Drepanosiphinae: Phyllaphidini

Both the known species live on Magnolia in East Asia. They are reviewed by
Takahashi (1965b), Higuchi (1972) and Moritsu (1983; including colour photo-
graphs of both species). Sexuales and life cycle are not recorded for either
species.
Neocalaphis magnoliae (Essig and Kuwana) All adult viviparae alate, pale
shiny straw yellow to yellow-green with antennae and bases of tibiae banded
with black, and forewing veins conspicuously black-bordered; BL 1.4-2.2 mm.
On leaves of Magnolia kobus and M. liliflora in Japan and Korea. 2n = 20
(Blackman, 1986).
Neocalaphis magnolicolens (Takahashi) All adult viviparae alate, almost
translucent yellowish-white to very pale green; BL 2.3-3.3 mm. On undersides
of leaves of Magnolia obovata in Japan. Immatures feed alongside main veins.
2n = 20 (Blackman, 1986).

NEOCHROMAPHIS Takahashi Drepanosiphinae: Phyllaphidini

Two East Asian species feeding on twigs and branches of Betulaceae. They
are stout-bodied aphids with pigmented wings, long dorsal abdominal hairs
and a very short processus terminalis. Higuchi (1972) reviewed the Japanese
species. (A third, western palaearctic species on Ostrya, ostryae Börner, was
placed in this genus by Eastop and Hille Ris Lambers (1976), but is here
regarded as a Pterocallis.)
Neochromaphis carpinicola (Takahashi) All viviparae alate, stout-bodied,
dark brown with forewings almost wholly pigmented (see Moritsu, 1983);
BL 1.7-2.0 mm. On undersides of twigs and branches of Carpinus spp. in
Japan and Korea. Alate males and apterous oviparae in October-November
(Takahashi, 1923).
774 Neohormaphis

Neochromaphis coryli Takahashi All viviparae alate, stout-bodied, bluish-

green or brownish, slightly shiny, with maculate forewings, the veins ending
in dark spots; BL 1.6-2.2 mm. On young shoots, twigs and branches of
Corylus spp. in China, Japan, Korea and Siberia. Also recorded from
Carpinus laxiflora (Shinji, 1941). Sexuales and life cycle not recorded. 2n = 18
(Chen and Zhang, 1985b).

NEOCRANAPHIS Ghosh and Quednau

Drepanosiphinae: Phyllaphidini

One or two Asian bamboo-feeding species originally placed in Shivaphis,

but distinguished by the setiform empodial hairs and other characters (A.K.
Ghosh and Quednau, 1990). Cranaphis and Phyllaphoides are also closely
related.

Neocranaphis arundinariae (Takahashi) All viviparae alate, elongate-bodied,

white, slightly dusky in thorax but without dorsal markings, secreting cottony
wax on anterior part of body, hind legs and (slightly) on antennae; BL
c. 2.1 mm. On undersides of leaves of Arundinaria (Pleioblastus) sp. in Taiwan
(Liao, 1976; as Shivaphis arundinariae). Sexual morphs and life cycle unknown.

Neocranaphis bambusicola David, Rajasingh and Narayanan All viviparae

alate, elongate-bodied, white with powdery wax covering body, legs and
antennae: BL 2.7-3.0 mm. On undersides of leaves of Bambusa sp. in West
Bengal, India. An ovipara and an apterous oviparoid (intermediate) female
were collected in November (David et al., 1970). The differences between this
species and N. arundinariae, apart from size, are unclear and its separate iden-
tity requires confirmation.

NEOHORMAPHIS Noordam Hormaphidinae: Nipponaphidini

One species in Java with unusual wing venation.

Neohormaphis calva Noordam Galls on upper sides of leaves of Distylium

stellare in Java are pale greenish, semiglobose, downy, about 1.5 cm in
diameter, usually arising singly from mid-rib or a smaller vein. Alatae (BL
1.4-1.8 mm, with black eyes and dark grey abdomen) leave gall in September
through a small hole on the underside of the leaf (Noordam, 1991). Migration
occurs to Quercus sp.; BL of apterae on Quercus is 0.8-1.0 mm, appearance
in life unknown, probably covered with wax powder.
 Neonipponaphis 775

NEONIPPONAPHIS Takahashi Hormaphidinae: Nipponaphidini

A genus for one species with prosoma distinctly separate from ABD TERG
2-7 and bearing numerous fine hairs.

Neonipponaphis shiiae Takahashi Apterae blackish-brown to almost black

in life, but pale in cleared preparations; BL 1.3-1.7 mm. On branches and
shoots of Castanopsis cuspidata in Japan (Takahashi, 1962a). Alatae are
black with dusky wings. Life cycle is unknown.

NEOPEMPHIGUS Mamontova and Kolomoets

Pemphiginae: Pemphigini

One palaearctic species feeding on poplar roots, closely related to Pachypappa

and possibly belonging in that genus.

Neopemphigus turajevi Mamontova and Kolomoets In dense wax-dusted

colonies causing gall-like swellings on fine roots of Populus bolleana (= P.
alba var. pyramidalis) in Ukraine (Mamontova and Kolomoets, 1981). Alate
sexuparae appear in September. Apparently monoecious on Populus, but
the life cycle needs to be clarified. Severe damage by this aphid is reported
(Kolomoets, 1984).

NEOPHYLLAPHIS Takahashi Drepanosiphinae: Phyllaphidini

About 12 species on Podocarpaceae and Araucariaceae in the southern

hemisphere and mountains of the tropics, extending northward into China and
Japan. An unusual feature of Neophyllaphis is that several species have
winged oviparae. Where sexual morphs are known, they generally occur in
mixed colonies with parthenogenetic morphs, often over several generations
in spring, summer or autumn. Hille Ris Lambers (1967) and Russell (1982)
provided reviews. The structure and function of adhesive vesicles on the
posterior abdominal sternites was studied by White and Carver (1971).

Neophyllaphis araucariae Takahashi Apterae broadly oval, bright yellow to

yellowish-orange; BL 1.1-1.5 mm. On young growth of Araucaria spp. in
776 Neophyllaphis

Mauritius (original description), Java, New Guinea, Australia, Hawaii, Costa

Rica (BMNH colln), Mexico and USA (Florida, California). Perhaps native
to Norfolk Island or Australia (Carver, 1959), or Southeast Asia. Probably
mainly anholocyclic, but the alate ovipara is known from Hawaii (Carver,
1971). 2n = 18 (Hales and Lardner, 1988).

Neophyllaphis brimblecombei Carver Apterae cream-coloured under a

coating of greyish wax; BL 1.4-2.1 mm. On Podocarpus elatus in Queensland,
Australia. Specimens from Podocarpus chinensis in Hong Kong (BMNH
colln) also key to this species. Hales (1976) gave an account of the biology
(although her aphids on P. spinulosus were N. lanata). Alate oviparae and
males occur in spring and summer (Carver, 1971); eggs and oviposition were
described by Carver and Hales (1974). 2n = 26 (Hales and Lardner, 1988).

Neophyllaphis fransseni Hille Ris Lambers Appearance in life not recorded;

BL of aptera 1.0-1.6 mm. On Podocarpus spp. in Java. Alate oviparae are
produced in January-March (Russell, 1982; and BMNH colln, leg. DHRL).

Neophyllaphis gingerensis Carver Apterae purplish-blue, with a coat of

white wax; BL 1.4-1.8 mm. On Podocarpus alpina, living singly or in small
groups on leaves, in the mountains of Australian Capital Territory. Apterous
oviparae and alate males in January-February (Carver, 1959). This is the only
species of the genus known to have apterous oviparae, perhaps as an adapta-
tion to the mountain habitat (Eastop, 1966). Milner and Holdom (1986)
recorded a fungal pathogen. 2n = 14 (Hales and Lardner, 1988).

Neophyllaphis grobleri Eastop Apterae are orange-coloured under a thick

coat of wax; BL 1.7-1.8 mm. On Podocarpus spp. in Africa (Angola,
Cameroon, Ethiopia, Kenya, South Africa, Tanzania), feeding on both young
and full-grown trees. Heavy attack can stunt growth of seedlings (Eastop,
1955a). Alate oviparae (for description see Russell, 1982) and males (Dürr,
1971) in spring. Ponsen (1990) studied the digestive system. 2n = 18*.

Neophyllaphis lanata Hales and Lardner Apterae deep reddish-brown to

reddish-purple, covered in a woolly white wax exudate; BL c. 1.5mm. On
stems and leaves of young shoots of Podocarpus spinulosus (a shrub). Hales
and Lardner (1988) described enzyme, chromosome and other differences
from N. brimblecombei, with which it was previously confused. Alate oviparae
and males start to appear in early spring (September). 2n = 24 (Hales and
Lardner, 1988).

Neophyllaphis michelbacheri (Essig) Apterae broadly oval, whitish-grey in

life due to covering of wax powder; BL c. 1.8mm. Alatae have dark dorsal
abdominal cross-bands. In Chile. Described from Pilgerodendron uviferum,
but also recorded (Carrillo, 1980) from Podocarpus nubigena, which is more
likely to be the normal host. Alate oviparae and males in January (Russell,
1982). Oviparae are unusual in having numerous scent plaques on fore and
middle tibiae as well as on the hind tibiae (Essig, 1953).
 Neoprociphilus 777

Neophyllaphis podocarpi Takahashi Apterae broadly oval, reddish-purple

thickly dusted with bluish-white wax; BL 1.5-2.0 mm. On leaves, young twigs
and fruit stems of Podocarpus spp. in China, Japan, Taiwan, Java, Malaya,
and introduced to North America and Hawaii. It causes stunting and curling
of new growth (Johnson and Lyon, 1988, p. 294). Alate oviparae and males
in both spring and autumn in Japan (Shibata, 1955); elsewhere, sexual morphs
appear rarely. Russell (1982) summarizes all available information. 2n = 26
from Japan (Blackman, 1986), or 2n = 24 from China (Chen and Zhang,
1985b); perhaps indicative of more than one species under this name.

Neophyllaphis podocarpini Carrillo, Plate 8 c, d Green and red in life;

BL c. 2.2 mm. On Podocarpus salignus in Chile; this species could not live on
P. nubigena (Carrillo, 1980). Alate oviparae and males occur along with
parthenogenetic morphs in December.

Neophyllaphis rappardi Hille Ris Lambers Apterae dark brown-purple,

covered with wax; BL c. 1.7 mm. On Agathis labillardieri in Biak, Irian Barat.
Found in small colonies on undersides of very young needles of older trees
in deep forest shade (Hille Ris Lambers, 1967). Sexual, morphs unknown.

Neophyllaphis totarae Cottier Apterae dark purplish-brown, dusted with

whitish wax powder; BL c. 2.8mm. On Podocarpus spp. in New Zealand,
usually feeding singly on leaves or in small colonies on young twigs. Alate
oviparae and males in spring and summer (Cottier, 1953, and BMNH colln).
2n = 10*.

Neophyllaphis viridis Ilharco Apterae dark green, covered with thick wax
powder; BL 1.5-1.8 mm. On Podocarpus falcata in southern Africa, found
in a mixed colony with N. grobleri (Ilharco, 1973; Dürr and Martin, 1976).
The alate vivipara was described by Russell (1982). Sexual morphs unknown.

NEOPROCIPHILUS Patch Pemphiginae: Pemphigini

One nearctic species with host alternation between Acer and Smilax.

Neoprociphilus aceris (Monell) Apterae in life covered in white wax; BL

3.3-5.3 mm. On undersides of twigs and branches of Acer saccharum in
spring, sometimes extending onto petioles and leaves. Throughout range of
A. saccharum in eastern North America. Heteroecious holocyclic; alatae
produced in May-June migrate to Smilax spp. (Smith and Graham, 1967).
Sexuparae return to A. saccharum in late August-October, although over-
wintering can also occur as hibernating close-knit clusters of first instar
nymphs in leaf litter, even in regions with severe winters outside the range
of A. saccharum (Lee and Robinson, 1978). 2n = 14.
778 Neosensoriaphis

NEOPTEROCOMMA Hille Ris Lambers

Aphidinae: Pterocommatini

A little-known genus of three species, two from basal and underground parts
of Salix in Europe, the third from Populus in China. No alate morph is
known, and in the one case where sexuales have been collected the males
appear to be paedogenetic.

Neopterocomma asiphum Hille Ris Lambers Apterae reddish (Heinze,

1961b), probably wax-covered; BL 2.5-3.1 mm. On lower part of trunk of
Salix spp. (alba, fragilis), attended by ants. Only recorded from Germany.
Other morphs and life cycle unknown.

Neopterocomma populivorum Zhang Appearance in life not recorded; BL

of aptera c. 2.8 mm. On Populus sp. (feeding site not recorded) on Changbei
Mountain, Jilin, China (Zhang and Zhong, 1990). Other morphs and life cycle
unknown.

Neopterocomma verhoeveni Hille Ris Lambers Apterae covered with fine

wax powder, dirty reddish or purplish-grey; BL 2.0-2.6 mm. In small colonies
on underground parts of older branches of Salix repens, attended by ants.
Only known from The Netherlands. Monoecious holocyclic; sexuales in
October. Males are yellowish-white, dwarfish and apparently only moult once
to attain maturity (Hille Ris Lambers, 1956b), although this is so unusual as
to require further confirmation.

NEOSENSORIAPHIS Quednau Drepanosiphinae: Phyllaphidini

A genus for one Nothofagus-feeder related to Neuquenaphis, but the aptera

has finger-like processes only on ABD TERG 7 and 8 and a strong pattern
of polygonal reticulation of the dorsal cuticle (Quednau, 1990).

Neosensoriaphis parva Quednau Appearance in life unknown, probably

dark under a covering or dusting of wax; BL of aptera 1.0-1.4 mm. On
Nothofagus obliqua (incl. var. macrocarpa) in Chile. Biology unknown;
an apparent fundatrix and a single alata were found in November (BMNH
colln, leg. DHRL).
 Neosymydobius 779

NEOSYMYDOBIUS Baker Drepanosiphinae: Phyllaphidini

About ten species resembling Symydobius but associated with nearctic oaks.
They usually feed on bark of terminal twigs, extending onto leaf petioles and
along basal parts of main veins. Remaudière (1982a) provided a key.

Neosymydobius agrifoliae (Essig) Apterae plump-bodied, dark reddish-

brown with a paler yellowish Y-shaped mark on dorsum (Essig, 1917), banded
antennae and dark hind legs; BL 1.6-2.3 mm. Alatae have dark-bordered
wing veins. Monoecious holocyclic on Quercus spp. (agrifolia, chrysolepis,
douglasii) in California. Apterous males and oviparae in October (BMNH
colln, leg. R.C. Dickson).

Neosymydobius ajuscanus Remaudière Apterae somewhat flattened dor-

soventrally, yellowish-white with black-banded antennae; BL 1.1-1.8 mm.
Collected on undersides of leaves of Quercus mexicana, and from Q. grisea,
in Mexico. Alatae present in September-October. Biology and sexuales
unknown.

Neosymydobius albasiphus (Davis) Apterae and alatae shining black or

brown with a whitish spinal stripe (Hottes, 1926) and/or white waxy areas
around pale siphunculi (Richards, 1968d), banded antennae and black hind
legs; BL 1.4-1.9 mm. On Quercus spp. (alba, macrocarpa, michauxii,palustris,
stellata) in eastern North America. Oviparae and alate males in September-
October (Davis, 1914 and BMNH colln, leg. RLB). Neosymydobius cana-
densis Richards, 1965, described from Q. macrocarpa in Ontario, differs only
in absence of the white waxy areas around the siphunculi and possibly in
larval chaetotaxy; it was synonymized with albasiphus by Remaudi re (1982a).

Neosymydobius butzei Remaudière Apterae dark shiny chocolate brown,

with black hind legs; BL 1.6-2.1mm. On Quercus mexicana in Mexico;
Remaudière (1982a) also described a variant form (Var. papillata'), which was
found on Q. centralis and Q. peduncularis as well as Q. mexicana. Biology
and sexuales unknown; believed to be anholocyclic.

Neosymydobius chrysolepis Swain Apterae and alatae brown, with conspi-

cuously pale siphunculi; BL 1.7-2.1mm. On Quercus spp. (chrysolepis,
engelmanni, gambelii) in western USA (California, Utah). Oviparae and alate
males in October (BMNH colln, leg. G.F. Knowlton).

Neosymydobius luteus Tissot Aptera yellowish (only one aptera is known,

and they are apparently rare in comparison with alatae). Alatae have brown
head and thorax and pale green-yellow abdomen; BL 1.2-1.4 mm. On Quercus
780 Neothoracaphis

nigra and Q. cinerea in Florida (Tissot, 1932a) and Louisiana. Biology and
sexuales unknown.

Neosymydobius memorialis Hottes and Frison Apterae with yellowish head

and abdomen and pale brown thorax; BL 1.2-1.5 mm. Alatae have chestnut-
brown head and thorax and lemon-yellow abdomen. On Quercus stellata in
Illinois, USA and also recorded from Q. lyrata in Louisiana (BMNH colln,
leg. H.B. Boudreaux). Oviparae in October in Illinois (Hottes and Frison,
1931).

Neosymydobius mimicus Hottes Apterae shining dusky to dark greenish-

brown; BL c. 1.6mm. Alatae have a green abdomen with darker brownish-
green cross-bars. On Quercus alba in Minnesota, USA (Hottes, 1926) and also
recorded from Q. macrocarpa in Manitoba (Robinson, 1971). Biology and
sexuales unknown.

Neosymydobius quercihabitus Miller Apterae and alatae dark brown, with

conspicuously banded antennae; BL 1.5-2.2 mm. On Quercus stellata in
eastern USA (Maryland, Massachussetts). Quednau (1971) provided a rede-
scription. Biology and sexuales unknown.

NEOTHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini

About ten species mainly known from Quercus in East Asia. Apterae are
black, sclerotic, lack siphuncular pores and have much reduced tarsi, with the
fore and mid-legs concealed but the hind legs usually projecting a little behind
the body. Host alternation and gall formation on Distylium is only established
for one species (yanonis); several species seem to be anholocyclic on evergreen
Quercus, with the apterous vivipara the only known morph. Most species were
originally described in Thoracaphis. Takahashi (1958d) and A.K. Ghosh and
Raychaudhuri (1973a) gave accounts of the genus and keyed species under
Microthoracaphis, which is here included within Neothoracaphis. Tao (1966)
keyed the species in Taiwan.

Neothoracaphis elongata (Takahashi) Apterae elongate oval, black, with

pale hind legs projecting behind body; BL c. 0.7 mm. Living throughout the
year on foliage of evergreen Quercus spp. in Japan (Takahashi, 1958d). (N.B.
not to be confused with Parathoracaphis elongata (Takahashi), which is a
quite different species.)

Neothoracaphis garhwalensis Chakrabarti and Raha Apterae oval, black,

with pale hind legs projecting a little behind body; BL 0.7-0.8 mm. In
 Neothoracaphis 781

ant-attended colonies on leaves of Quercus spp. (incana, serrata) in Uttar

Pradesh, India (Chakrabarti and Raha, 1985). Life cycle unknown.

Neothoracaphis glaucae (Takahashi) Apterae oval, indented at frontal

margin, black, with much wax mid-dorsally and yellow hind legs protruding
behind body; BL 0.8-1.0 mm. Living throughout the year on foliage of
Quercus glauca in Japan (Takahashi, 1958d).

Neothoracaphis quercicola (Takahashi) Adult apterae black, broadly oval,

without wax; BL c. 0.9mm (Takahashi, 1924a; as Astegopteryx). Immatures
are yellow and secrete much wax. Alatae have black head and thorax and
dusky yellowish-brown abdomen. Living parthenogenetically all year on
undersides of leaves of Quercus variabilis (records from Q. serrata are
probably in error) in Taiwan. Alatae are produced in November-March
(Takahashi, 1931a).

Neothoracaphis querciphaga (Takahashi) Apterae black, narrowly oval,

with protruding hind legs; BL c. 0.8 mm. Living all year on foliage of Quercus
myrsinaefolia in Japan (Takahashi, 1958d).

Neothoracaphis saramaoensis (Takahashi) Apterae black, elongate oval,

indented on frontal margin, with white wax dorsally, including a pair of long
whitish wax 'horns' arising from middle of anterior part of prosoma, and also
'a wide, very thin whitish secretion behind the hind end' (Takahashi, 1935c).
Hind legs projecting posteriorly, pale brown. BL 0.5-0.7 mm. On undersides
of leaves of Quercus variabilis in Taiwan and populations on Q. glauca in
Japan are also thought to be this species (Takahashi, 1958d). Life cycle
unknown.

Neothoracaphis sutepensis (Takahashi) Apterae broadly oval, black with

much white wax mid-dorsally and black hind legs protruding behind body;
BL c. 0.6 mm. On undersides of leaves of a tree of the Fagaceae (probably
Quercus sp.) in Thailand (Takahashi, 1941). Life cycle unknown.

Neothoracaphis tarakoensis (Takahashi) Apterae broadly oval with frontal

margin indented, black, with much white wax mid-dorsally and black hind legs
protruding behind body; BL 0.9-1.0 mm. On undersides of leaves of Quercus
spinosa in Taiwan (Takahashi, 1937b). Life cycle unknown. The distinction
between this species and N. yanonis, apart from size, is unclear.

Neothoracaphis yanonis (Matsumura) Host-alternating between Distylium

racemosum and deciduous Quercus spp. (dentata, serrata) in Japan, Korea
and China. Small, prominent galls develop in April-May, protruding from
both sides of Distylium leaves (Fig. 130B, p. 785; and Moritsu, 1983, as Nip-
ponaphis yanonis). All second generation are alatae, with black head, thorax,
antennae and legs, yellowish-brown abdomen and dark-bordered wing veins;
BL 1.2-1.6 mm. Apterae on undersides of Quercus leaves are black with
782 Neuquenaphis

dorsal wax and black hind legs projecting behind body; BL 0.5-0.6 mm. Alate
sexuparae return to Distylium in October-November and produce abundant
sexuales on undersides of leaves (Takahashi, 1958d). Shen (1986) studied the
life cycle and chemical control measures in China and Ito (1984) recorded
invasion of galls by the kleptoparasitic moth Nola innocua in Japan. Neo-
thoracaphis hangzhouensis Zhang, described from China (Zhang and Zhong,
1982b), is not clearly distinct from N. yanonis and may be a synonym.

NEOTUBERAPHIS Pal and Raychaudhuri

Hormaphidinae: Cerataphidini

Two species in India, one on Eugenia and the other described from an
unnamed parasitic plant on a tree (Pal and Raychaudhuri in Raychaudhuri
et al., 1980c). Raha and Raychaudhuri (1981) provided a key.

Neotuberaphis indica Raha and Raychaudhuri Apterae are dark brown, BL

1.3-1.5 mm, living in white wax in marginal leaf-folds on Eugenia sp. in
Nagaland, India. Alatae have pale brown abdomen and dark wing veins
(Raha and Raychaudhuri, 1981).

NEUQUENAPHIS Blanchard Drepanosiphinae: Phyllaphidini

A South American genus of about ten species associated with Nothofagus,

closely related to Taiwanaphis, in East Asia, Australia and New Zealand. Hille
Ris Lambers (1968) revised the genus. Their biology is little known; for some
species only the aptera or the alata is described. Several species seem to feed
on more than one species of Nothofagus, but taxonomy and host relations
need further investigation.

Neuquenaphis bulbicauda Hille Ris Lambers Colour in life unknown, but

probably rather dark with a covering of powdery wax; BL of apterae 2.2-
3.1 mm. On leaves of Nothofagus dombeyi in Chile (Hille Ris Lambers, 1968).
[The alata and sexual morphs described under bulbicauda are other species.
The alata is Neuquenaphis valdiviana Carrillo (Quednau, pers. comm.). Eastop
and Hille Ris Lambers (1976) synonymized bulbicauda with Neuqenaphis
chilensis Essig, but Quednau (pers. comm.) points out that this is not tenable;
alatae with a short ANT PT and dark-bordered wing veins collected from
Nothofagus dombeyi are N. chilensis. Apterae and sexual morphs of another,
undescribed, species have also been collected on Nothofagus dombeyi; this
 Neuquenaphis 783

species is similar to bulbicauda but the apterae have only 4 processes on ABD
TERG 7 and few ventral cephalic hairs (Quednau, pers. comm.).]

Neuquenaphis edwardsi Laing ( = flavipes Hille Ris Lambers, = papillata

Richards) Colour in life unknown, probably rather dark with a covering of
powdery wax; BL of apterae and alatae 1.8-2.6 mm. On various Nothofagus
spp. in Argentina and Chile, including Tierra del Fuego. Apterous oviparae
and alate males in April-May (BMNH colln, leg. G. Monsalve).

Neuquenaphis essigi Hille Ris Lambers Appearance in life unknown, pro-

bably rather dark, perhaps with greyish wax. Apterae very broadly oval; BL
of apterae and alatae 2.0-2.4 mm. On Nothofagus obliqua (incl. var. macro-
carpa) in Chile (Hille Ris Lambers, 1968). Sexuales unknown.

Neuquenaphis michelbacheri Essig Appearance in life unknown, probably

greyish due to covering of wax powder; BL of alata 1.7-2.3 mm. Apterae
undescribed and possibly not produced. Very similar to N. edwardsi, which
apparently feeds on a similar range of Nothofagus species, but can be dis-
tinguished by embryonic chaetotaxy (F.W. Quednau, pers. comm.).

Neuquenaphis palliceps Hille Ris Lambers Appearance in life unknown;

larger and probably paler than most other described species; BL of apterae
and alatae 2.2-2.9mm. On Nothofagus spp., especially N. dombeyi, in
southern Argentina and southern Chile. Apterae collected in November-
December were possibly fundatrices (Hille Ris Lambers, 1968), and an alate
male was collected in April (BMNH colln, leg. G. Monsalve).

Neuquenaphis schlingeri Hille Ris Lambers Appearance in life unknown,

probably rather pale; BL of apterae and alatae 1.4-1.8 mm. On Nothofagus
obliqua in Chile (Hille Ris Lambers, 1968). Alate males and apterous oviparae
in April-May (BMNH colln, leg. G. Monsalve).

Neuquenaphis sensoriata Hille Ris Lambers, Plate 8a, b Appearance of

adults in life unknown, probably rather dark with dark abdominal cross-bands
and a dusting of wax; BL of apterae and alatae 1.7-2.1 mm. Immatures are
shiny red-brown (slide label, BMNH colln). On Nothofagus spp. (especially
glauca, obliqua) in Chile. Fundatrices in November (BMNH colln, leg.
DHRL), sexuales unrecorded.

Neuquenaphis similis Hille Ris Lambers Apterae elongate oval, of unknown

colour but probably rather pale; BL 2.6-2.8 mm. Alatae are unknown. In
Chile; described from Nothofagus obliqua (Hille Ris Lambers, 1968), but all
subsequent collections, including possible fundatrices in November, were
from N. pumilio (BMNH colln, leg. DHRL).
784 Nipponaphis

NIPPOLACHNUS Matsumura Lachninae: Lachnini

An oriental genus of four or five species characterized by the absence of an

ocular tubercle (triommatidium). Associated mainly with Eriobotrya, but
parthenogenetic generations may utilize other woody Rosaceae. A.K. Ghosh
(1982b) reviewed the genus.

Nippolachnus bengalensis Basu and Hille Ris Lambers Apterae long-bodied,

pale green; BL 2.6-3.3 mm. Alatae have a variable sclerotic pattern on the
dorsal abdomen. On undersides of leaves of Eriobotrya dubia in West Bengal,
India (Basu and Hille Ris Lambers, 1968); also recorded from Pyrus pashia
and Photinia arguta in Meghalaya and from cultivated pears in Bhutan
(Agarwala, 1983). Specimens from Pyrus granulosa in Malaya (Takahashi,
1950; as N. piri) appear to be this species. Life cycle and sexual morphs are
unknown.

Nippolachnus himalayensis (van der Goot) Apterae broadly oval, dirty

greyish to brick red with whitish intersegmental bands of wax and mainly
black legs: BL 3.5-5.5 mm. In large colonies on undersides of leaves of
Eriobotrya petiolata in West Bengal, India (as eriobotryae; A.N. Basu and
Hille Ris Lambers, 1968). Life cycle and sexual morphs unknown.

Nippolachnus piri Matsumura, Plate 15a, b Apterae long-bodied, pale green

with some bluish-white dorsal wax, with pale brown legs and antennae; BL
3.0-3.7 mm. On undersides of leaves, feeding along main veins, of Eriobotrya
japonica and also recorded from Pyrus spp., Raphiolepis umbellata and
Sorbus alnifolia. In China, India, Japan, Korea and Taiwan. Oviparae and
alate males occur on E. japonica in Japan in October-November (Takahashi,
1923). In India (West Bengal, Sikkim) populations are apparently anholocyclic
and host-specific to Pyrus communis, on which large damaging populations
may build in the summer months (M.R. Ghosh and Raychaudhuri, 1981).
Nippolachnus xitianmushanus, on E. japonica in China, may or may not be
distinct (Zhang and Zhong, 1982b).

NIPPONAPHIS Pergande Hormaphidinae: Nipponaphidini

About 15 East Asian species, some with host alternation between galls
on Distylium and Fagaceae, others apparently anholocyclic on Fagaceae,
Lauraceae and Moraceae. Apterae on secondary hosts are aleyrodiform, with
the strongly sclerotized prosoma bearing pustulate ornamentation. Alatae
 Nipponaphis 785

have smoky wings. Takahashi (1962a) reviewed the species on Distylium in

Japan and Sorin (1987b) illustrated differences in embryonic chaetotaxy.
Many of the Japanese species are illustrated by Moritsu (1983). Noordam
(1991) revised the species in Java. A.K. Ghosh and Raychaudhuri (1973b)
reviewed and keyed the genus; the three species they placed in subgenus
Pseudonipponaphis are here transferred to Schizoneuraphis.
Nipponaphis brevipilosa Noordam Apterae pale brown with antennae, legs
and margins of abdominal plate yellowish: BL 1.3-1.6 mm. In dense colonies
on young and older twigs of Castanopsis argentea in Java (Noordam, 1991).
Alatae were collected in May. Life cycle unknown.

Nipponaphis distychii Pergande Large galls on stems of Distylium race-

mosum in Japan and Korea purported to be formed by N. distychii are

Fig. 130. Galls of Nipponaphidini on Distylium. A, Dinipponaphis autumna (after Moritsu, 1983);
B, Neothoracaphis yanonis (after Paik, 1972); C, Monzenia globuli (after Moritsu, 1983);
D, Metanipponaphis cuspidatae (after Sorin, 1987b); E-G, (3 figs) various galls of Nipponaphis monzeni
(all from Sorin, 1958); H, N. litseae (after Sorin, 1987b); I, N. ?machili (after Sorin, 1987b);
J, Schizoneuraphis gallarum (after Docters van Leeuwen-Reijnvaan and Docters van Leeuwen, 1926).
A-l, all on Distylium racemosum; J, on D. stellare.
786 Nipponaphis

illustrated by Moritsu (1983) and Paik (1972) respectively; however, there is

some doubt about the species causing these galls, as the original description
was of alatae emerging from galls on leaves. Alatae emerge in autumn;
according to the original description (Pergande, 1906, quoting Kuwana)
they have black head and pterothorax, purplish-yellow prothorax and dark
purplish-brown abdomen; BL 1.9-2.2 mm. Antennal rhinaria are distributed
III 39-45, IV 21-25, V 10-13. The wings are only slightly dusky - more so
at their bases. The secondary host is unknown. Mpnzen's (1954) account
under the name distychii refers to Sinonipponaphis monzeni (Takahashi,
1962a).

Nipponaphis distyliicola Monzen, Plate 3e, f Oblong galls are produced on

the twigs of Distylium racemosum in Japan (Monzen, 1934). Alatae (BL
1.5-2.5 mm, antennal rhinaria distributed III 21-39, IV 10-19, V 8-14) emerge
in autumn and fly to Quercus spp. (acuta, glauca, crispula). Records from
Castanopsis cuspidata are probably based on misidentifications. Apterae on
branches and shoots of Quercus are almost circular, black; BL 1.2-2.2 mm.
Alate sexuparae (BL 2.5-2.7 mm) return to Distylium in April, but anholo-
cyclic populations also persist on Quercus (Takahashi, 1962a; Sorin, 1958, as
gigantea). Kurosu et al. (1990) described 'soldiers' in the galls and Ito and
Hattori (1982) reported predation by Nola innocua. Thoracaphis coreana on
Quercus myrsinaefolia in Korea (Paik, 1965) appears to be a synonym, to
judge from paratypes in the BMNH collection.

Nipponaphis ficicola Hille Ris Lambers and Takahashi Apterae very broadly
oval, wrinkled, dark purple to blackish in life; BL 1.0-1.3 mm. On the bark
of about 1 cm diameter branches and green twigs of Ficus benjamina and Ficus
sp. in Java (Hille Ris Lambers and Takahashi, 1959). Apterae collected in
June-November. Life cycle unknown.

Nipponaphis javanica Noordam Apterae brownish, with some wax powder:

BL c. 1.8 mm. In dense colonies on young shoots, and on under and upper
sides close to mid-ribs of youngest leaves, of Castanopsis acuminatissima in
Java (Noordam, 1991). Alatae were collected in August and January. Life
cycle unknown.

Nipponaphis karatanei (Sasaki) This species, described from Magnolia

fuscata in Japan (Sasaki, 1936) was placed in Nipponaphis by Monzen (1954),
but is unlikely to belong to this genus. The association with Magnolia is also
doubtful, because the galls described appear to be those of a psyllid that lives
on Persea, and the description of the aphid has many inconsistent features
making its taxonomie position uncertain.

Nipponaphis litseae Takahashi Spindle-shaped, pale green galls, c. 5.6 cm

long when mature, are produced on twigs of Distylium racemosum in Japan
(Fig. 130 H; and Sorin, 1987b). Alatae (BL not recorded, secondary rhinaria
distributed III 43-46, IV 20-22, V 11-15) migrate to Litsea glauca and
 Nipponaphis 787

Neolitsea sericea, where the blackish-brown, almost circular apterae feed on

the branches (Takahashi, 1959b).

Nipponaphis machili (Takahashi) Galls on twigs of Distylium in Japan

believed to belong to this species are glossy, green, ovoid, c. 3.3 cm long when
mature (Fig. 130I), giving rise to alatae with antennal rhinaria distributed III
62-73, IV 28-31, V 20-24 (Sorin, 1987b). Migration occurs to leaves of Persea
thunbergii in Japan. The almost circular, blackish-brown apterae (BL c. 1 mm)
were originally described from the undersides of leaves of Machilus (= Persea)
in Taiwan (Takahashi, 1933b).

Nipponaphis machilicola (Shinji) Apterae almost circular, dorsally flattened,

dark bluish-brown with pale yellow markings (see Moritsu, 1983); BL 1.5-
2.2mm. On branches of Persea sp. in Japan. But for feeding site, body
size and greater variation in shape of dorsal pustules, this species strongly
resembles N. machili, and its taxonomie status requires confirmation. Nip-
ponaphis amamiana Takahashi, 1962a appears to be a synonym.

Nipponaphis manoji Ghosh and Raychaudhuri Apterae almost circular,

dorsally flattened, dark brown, not strongly sclerotized; BL 1.2-1.4 mm
(A.K. Ghosh and Raychaudhuri, 1973b). In large, ant-attended colonies on
leaves and young stems of Litsea spp. (corymbosa, polyantha) in West Bengal,
India. Life cycle unknown; alatae (sexuparae?) were collected in February.
A Nipponaphis species described (but not named) from apterae on Litsea
polyantha in Nagaland (Raha and Raychaudhuri, 1981) is very similar, except
for the apparent lack of posteriomedial hairs on ABD TERG 7.

Nipponaphis monzeni Takahashi Galls on Distylium racemosum in Japan

are large, variously shaped, often pyriform or globular, pale brown, maxi-
mum diameter 2-6 cm (Figs 130E-G; Sorin, 1958; Moritsu, 1983). Alatae BL
c. 3.0mm) emerge from a round hole in the side of the gall in autumn and
migrate to Quercus glabra, where blackish subcircular apterae (BL 1.5-
1.7mm) live on the branches. Sexuparae return to Distylium in April. Eggs
hatch in May (Takahashi, 1962a).

Nipponaphis multisetosa Noordam Apterae brownish: BL 1.3-1.7 mm. On

young and older twigs of Castanopsis javanica in Java (Noordam, 1991).
Other morphs and life cycle unknown.

Nipponaphis semiglabra Noordam Aptera brown with a transparent coat of

wax, the prosoma having a paler brown spinal band, 2 paler thin transverse
lines along posterior margin of pronotum and ABD TERG 1, and 7 pairs
of small black spots: BL 1.2-1.5 mm. In small numbers on older twigs of
Lithocarpus bennettii in Java (Noordam, 1991). Other morphs and life cycle
unknown.
788 Oestlundiella

NURUDEA Matsumura Pemphiginae: Fordini

About five species forming galls on Rhus in East Asia and migrating in
autumn to unknown secondary hosts, likely to be mosses. Tsai and Tang
(1946) provided a review of the Chinese species and Tao (1970) keyed the
species in Taiwan (although his synonymy is not followed here). See also B
& E, 1984, pp. 188-189.

Nurudea ibofushi Matsumura (= sinica Tsai and Tang) Galls pale yellowish-
brown with a pinkish tinge, irregular in shape, max. diameter 4.5-5.5 cm; on
leaves of Rhus semialata in China and Japan. Alatae (BL c. 1 mm) with brown
head and thorax and pale yellow-green or yellow-brown abdomen emerge in
autumn (Tsai and Tang, 1946; as N. sinica).

Nurudea meitanensis (Tsai and Tang) Galls are rosy-red, somewhat flat-
tened, irregularly branched from the base, maximal dimension up to 10cm,
arising from leaflets of Rhus punjabensis var. sinica in China (Kweichow).
Alatae emerging from the gall in autumn have black head, pale green pro-
thorax, dusky pterothorax and pale yellowish abdomen (Tsai and Tang, 1946;
as Floraphis meitanensis). A closely-related species, N. choui Xiang, produc-
ing branched galls on Rh. potaninii in Shansi Province, differs in colour and
shape of gall, and the emergent alatae have longer ANT III and dense long
hairs on hind tibiae (Xiang, 1980/81).

Nurudea shiraii (Matsumura) Galls are densely pubescent, not particularly

reddish, on leaflets of Rhus semialata in China, Japan and Taiwan. Alatae
(BL 1.0-1.4 mm) with black head, blackish-brown pterothorax and yellowish-
brown abdomen, emerge in autumn and fly to mosses (Takahashi, 1943).

Nurudea yanoniella (Matsumura) (= Fushia rosea Matsumura) Galls are

rosy-red, irregularly branching outgrowths on leaflets of Rhus semialata in
China, Japan and Korea (Paik, 1972; as rosea). Alatae (BL 1.0-1.4 mm)
emerge in autumn and fly to an unknown secondary host.

OESTLUNDIELLA Granovsky Drepanosiphinae: Phyllaphidini

One North American species on Alnus, related to Euceraphis.

Oestlundiella flava (Davidson) All viviparae alate, delicate, lemon-yellow

with plumes of bluish-white wax developing on antennae and legs; BL
 Olegia 789

2.4-3.4mm. On undersides of leaves of Alnus spp., tending to aggregate on

certain leaves and feed along either side of mid-rib. Widely distributed in
North America (except the southeast). Apterous oviparae and alate males in
October. 2n = 8.

OLEGIA Shaposhnikov Phylloxeridae

One species with 1-segmented tarsi, forming galls on leaves of Japanese elm.

Olegia ulmifoliae (Aoki) Aptera (fundatrix?) in gall yellowish, broadly

pear-shaped; BL 1.0-1.7 mm. In small galls (Fig. 134A, p. 898; and Stoetzel,
1993) on the mid-ribs of leaves of Ulmus japonica in Japan (Hokkaido:
Aoki, 1973; as Aphanostigma) and eastern Siberia (Pashchenko, 1988b).
The fundatrix is remarkably fecund: Shaposhnikov (1979) counted over
1500 eggs and first instar larvae around one female in a gall in July. Alatae
are unknown, dispersal being by wind-blown first instars. Monoecious
holocyclic, sexuales mating either in the gall or on the leaf surface (Stoetzel,
1993).

OVATUS van der Goot Aphidinae: Macrosiphini

About 14 species of Myzws-like aphids, the alatae of which have no black

central abdominal patch. Palaearctic species are associated with Pyroidea
and/or Labiatae, whereas North American species are on a wide variety of
hosts. There are accounts for Europe (Heinze, 1960) and Japan (Miyazaki,
1971). (See also B & E, 1984, p. 321.)

Ovatus crataegarius (Walker) Apterae yellowish-green to mid-green; BL

1.4-1.9 mm. In spring colonies on undersides of young leaves of Pyroidea
(Crataegus, Cydonia or Malus), without causing leaf deformation. Hetero-
ecious holocyclic in Europe and North America, with Crataegus spp. as the
favoured primary hosts and summer generations on Labiatae (especially
Mentha). Anholocyclic on mints in warmer regions. In Japan, similar aphids
are recorded only from Cydonia and Malus and could be a separate taxon,
for which the name malicolens Hori is available. 2n = 12.

Ovatus insitus (Walker) Apterae greenish-white, rather shiny; BL 0.9-

1.9 mm. In spring on undersides of young leaves of Crataegus spp. or Mespilus
germanica, sometimes on Cydonia, Pyrus or Sorbus. Heteroecious holocyclic,
migrating for summer to Lycopus. In Europe and southwest Asia (Israel, Iran,
790 Pachypappa

Turkey). Müller and Hubert-Dahl (1979) compared it with the closely-related

O. crataegarius. 2n = 12.

Ovatus malisuctus (Matsumura) Apterae dark yellowish-brown to brownish-

green with black siphunculi and distal halves of femora; BL 1.0-1.7 mm.
Immatures shiny yellow (Moritsu, 1983). Curling young leaves of Malus spp.
(baccata, pumila, sieboldii) and Chaenomeles japonica. In China, Japan,
Korea, Taiwan and also recorded from Georgia (as Myzus chaenomelis
Dzhibladze). Monoecious holocyclic. Population ecology and control have
been studied in Japan and Korea (e.g. Takeda, 1979; Kim et al., 1986),
including interactions with predators (Hukusima, 1963). 2n = 12 (Chen and
Zhang, 1985b).

PACHYPAPPA Koch Pemphiginae: Pemphigini

About ten species mostly host-alternating between Populus and roots of Picea.
The fundatrices lack wax glands and the alate spring migrants have fore-
wings with once-branched media. Accounts are available from Heie (1980;
Fennoscandia and Denmark), Danielsson (1976, gall generations in Sweden;
and 1990a, root-feeding generations), Carter and Danielsson (1991, root-
feeding generations in Britain) and Smith (1974a; North America). Asiphum
is a synonym.

Pachypappa marsupialis Koch Forming large bag- or blister-like galls (Fig.

131E), open to the underside, on leaves of Populus nigra in Europe (Austria,
Estonia, Germany, Poland, Romania, Portugal). Fundatrix brownish-green
with white wax. Alatae leave the gall in June (Börner and Heinze, 1957); they
have a pinkish-yellow abdomen and BL 3.1-3.5 mm (Szelegiewicz, 1982). The
secondary host is unknown. A subspecies, P. marsupialis lambersi, produces
yellow blister-like galls on Populus spp. (laurifolia, maximowiczii) in Japan
and China (Aoki, 1976; Zhang et al., 1985). It has dimorphic first instar
larvae, one morph being migratory and leaving the gall to invade others (Aoki,
1979b). Alatae of ssp. lambersi migrate to an unknown secondary host in
July. 2n = 10 (for ssp. lambersi).

Pachypappa pilosa (Zhang) Appearance in life unknown: BL of emigrant

alata c. 3.2mm. On Populus sp. in Tibet (Zhang and Zhong, 1981b; as
Sigmacallis pilosa).

Pachypappa populi (Linnaeus) Forming large pale green, yellowish to

slightly reddish blister-like galls (Fig. 131F), open to the underside, on leaves
of Populus tremula in Europe (Fennoscandia, Latvia, Poland, the Alps) and
also reported (on P. tremula davidiana) from China (Zhang and Zhong,
 Pachypappa 791

1985b). Fundatrix bluish grey-black, with sparse or no wax; BL 5.1-6.3 mm.

Alatae, with olive green abdomen and BL c. 3.5mm, migrate in early July
in Europe, to found large colonies in flocculent wax on roots of Picea abies.
Sexuparae appear in September, but anholocycly on Picea roots also occurs
in southern Sweden (Danielsson, 1990a).

Pachypappa pseudobyrsa (Walsh) The fundatrix is wax-dusted, yellowish-

olive green with a blackish head; BL 4.0-4.7 mm Its feeding causes a small
blister-like swelling in the centre of the underside of the leaf near the mid-
rib. Its numerous progeny spread out in May-July along the main veins,
developing into dusky-dark alatae, BL c. 3.5 mm. On Populus spp. (deltoides,
fremontii, tremuloides) in eastern North America, where it is widely distri-
buted, and also in Colorado (Palmer, 1952, as Asiphum). Life cycle unknown;
the alatae presumably migrate to an unknown secondary host, although Smith
(1974a) thought that it might be monoecious.

Pachypappa rosettei (Maxson) Feeding by the fundatrix (BL 2.5-2.8 mm,

appearance in life unknown) causes bending of leaves and stunting of stems
to form a rosette-like nest, as in the closely-related European species
P. tremulae. On P. tremuloides in Colorado and Utah, USA. Alatae (BL 2.2-
2.5mm) are produced in July. Host alternation has not been experimentally
confirmed, but aphids on roots of Picea glauca, P. mariana and Pseudotsuga
menziesii have been tentatively assigned to P. rosettei (Danielsson, 1990a).

Pachypappa sacculi (Gillette) Fundatrix is wax-dusted, slate grey with

brown head; BL 4.5-5.0 mm. It forms a large bag-like gall on P. tremuloides,
with edges and tip of leaf curled downward (Harper, 1966; as Asiphum).
In western USA (Arizona, Colorado, Utah) and across Canada (British
Columbia, Alberta, Quebec, Ontario and Nova Scotia). Progeny of fundatrix
are all alate, with dark olive green abdomen, BL 3-4 mm. Host alternation
not experimenally confirmed, but aphids on roots of Picea spp. (pungens,
mariana) were tentatively assigned to P. sacculi (Danielsson, 1990a). 2n = 10.

Pachypappa shaposhnikovi Gabrid Forming large bag-like galls from leaves

of Populus spp. (densa, laurifolia) in Kirgizia (Gabrid, 1989). Alatae are very
large (BL c. 5 mm). Secondary host unknown.

Pachypappa tremulae (Linnaeus) Fundatrix on twigs of Populus tremula (or

more rarely P. x canescens) is almost globular, dirty reddish- or yellowish-
brown, without wax; BL 5.0-6.6 mm. Its progeny move onto new shoots and
cause bending of leaves and petioles and stunting of growth of shoot, forming
a rosette-like leaf nest (Fig. 131G). Second generation are all alate, orange-
yellow to pinkish-brown, wax-covered, with black head and pterothorax;
BL 3.2-4.4 mm. They migrate in June to found colonies on roots of Picea
abies (Stroyan, 1975). Apterous exules are pale yellowish-white with tufts of
wax posteriorly; BL 1.3-2.1mm. Sexuparae return to aspen in September-
October, but populations also overwinter anholocyclically on roots of Picea.
792 Pachypappella

Widely distributed in northern palaearctic, east to China (Zhang and Zhong,

1985b) and Japan (Aoki, 1976; as P. shirobamba). See Lavallee (1987) for a
bibliographic review. 2n = 10.

Pachypappa vesicalis Koch Forming irregular green to yellowish bag-like

or blister-like galls (see Danielsson and Carter, 1993b for photograph) on
leaves of Populus spp. (alba, canescens). Fundatrix is reddish-yellow to
orange-red, wax-dusted; BL 4.2-4.9 mm. All second generation are alate,
with reddish-brown abdomen, BL c. 3.3mm, migrating in June to found
colonies on roots of Picea spp. (abies, glauca, sitchensis). Apterous exules
on spruce roots are pale yellowish-white secreting white filamentous wax.
Sexuparae return to Populus in September-October; mating and oviposition
on poplar bark were described by Carter and Danielsson (1991). In northwest
Europe.

Pachypappa warshavensis (Nasonov) (= varsoviensis Mordvilko) Forming

loose leaf-nest galls on Populus alba and P. canescens (see Danielsson and
Carter, 1993b for photograph). Fundatrices (feeding on twigs) are red-brown;
BL 3.5-4.0 mm. Alatae yellowish-orange, BL 3.0-3.2 mm, migrating in June.
Apterae thought to be the secondary host generations of this species occur on
the roots of Salix caprea (Szelegiewicz, 1982, and BMNH collection), although
the transfer has not been accomplished experimentally (Danielsson, 1990a). In
Europe. 2n = 10*.

PACHYPAPPELLA Baker Pemphiginae: Pemphigini

One species closely related to Pachypappa, but the fundatrix has wax glands
on the thorax and abdomen.

Pachypappella lactea (Tullgren) The gall is formed on Populus tremula

by folding down of the leaf lamina to form a cone (Fig. 131H, p. 803), which
becomes bright orange. Fundatrices are greyish-brown with white wax; BL
c. 4 mm. All second generation are alate, with dark greyish-brown abdomen,
BL 2.5-3.5 mm, migrating in June-July to colonize roots of Picea abies
(Danielsson, 1990a). Anholocyclic overwintering on Picea roots, including P.
sitchensis (Carter and Danielsson, 1991), may be common. Northern palaearc-
tic and boreo-alpine in distribution, from Britain to Siberia (Pashchenko,
1988b).
 Paducia 793

PADUCIA Hottes and Frison Aphidinae: Pterocommatini

A genus of one or two little-known species. Its taxonomie position within

Aphidinae is uncertain.

Paducia antennata (Patch) Only oviparae (collected October-November)

and a single alate vivipara have been described (Patch, 1913a; Hottes and
Frison, 1931). Ovipara is dark greenish or chocolate brown to black dorsally,
greenish-yellow ventrally; BL c. 2mm. Alata mainly brown, BL c. 1.4mm.
Presumably monoecious holocyclic on roots of Salix sp., although the
evidence for this host association is entirely circumstantial. Recorded from
Illinois, New York and Maine, USA.

Paducia aterrima Hille Ris Lambers Apterae 'probably shiny black'; BL

2.0-2.2 mm. Other morphs not known. On roots of Salix sp. in Greenland
(Hille Ris Lambers, 1952). Some differences from P. antennata were given by
Hille Ris Lambers, but as different morphs were compared the distinctness
of the two species requires verification.

PANAPHIS Kirkaldy Drepanosiphinae: Phyllaphidini

Large distinctive aphids with short antennae living on Juglans regia (see B &
E, 1984, p. 248, as Callaphis). The three described taxa are closely related
and could perhaps be geographic variants of a single species. Previously the
name Callaphis Walker was widely used for this genus, but was suppressed
following a plea by Quednau (1983), on the grounds of potential confusion
with Calaphis Walsh.

Panaphis juglandis (Goetze) All viviparae alate, large and stout-bodied,

with dark head and thorax, yellow abdomen with dark brown transverse bands
and forewing veins fuscous-bordered; BL 3.5-4.3 mm. On upper sides of
leaves of Juglans regia, in rows along the veins, often ant-attended. In Europe,
Central Asia, Pakistan and northwest India and introduced into western USA.
Oviparae and alate males occur in September-October. Masoodi et al. (1987)
studied its population dynamics in Kashmir, India. Biological control has been
attempted in California with a Praon species from Afghanistan (Mackauer
and Sullivan, 1982). See also B & E, 1984, p. 248. 2n = 22*.

Panaphis nepalensis (Quednau) All viviparae alate, colour in life unrecorded

but probably much like P. juglandis (see Quednau, 1973). On Juglans regia
794 Paoliella

in Nepal and Tibet. Sexuales are unrecorded. Zhang in Zhang and Zhong
(1985c) described a subspecies, P. nepalensis yunnanensis, from Yunnan
Province, China, also collected on Juglans regia.

PAOLIELLA Theobald (= Unipterus Hall)

Drepanosiphinae: Phyllaphidini

About 22 species on Burseraceae and Combretaceae in Africa (plus one in

India), closely related to the South American genus Lizerius. Both apterous
and alate viviparae are known for most species, but there are a few species
which do not seem to produce apterae. Keys are provided by Ilharco (1971)
and Quednau (1974).

Paoliella ayari (Eastop) Apterae yellow, BL c. 0.9mm, on undersides of

leaves and young shoots of Commiphora spp. (pedunculata, angolensis).
Recorded from Nigeria, Tanzania, South Africa and Zimbabwe. Life cycle
unknown.

Paoliella browni Quednau The single known aptera, possibly a fundatrix,

was dark brownish-velvety-green with white dorsal processes; BL c. 1.1 mm
(Quednau, 1962; but the two alatae described in this paper are P. echinata).
On Combretum gueinzii (= C. molle) in South Africa. Possibly P. mono-
tuberculata, which forms dense colonies composed entirely of reddish-brown
alatoid nymphs on undersides of young leaves of Combretum sp. in Angola
(van Harten, 1979), is the same species, as suggested by Quednau (1974). Van
Harten (1979) described an aphid with greenish-yellow alatoid nymphs living
solitarily on undersides of very old leaves of Combretum sp., in Angola as
a subspecies, P. browni capinganae. Alate males and apterous oviparae of this
form (the latter dark velvety-green, BL 1.0-1.1 mm) were collected in early
June. An alata trapped in Burundi (Remaudière and Autrique, 1985) may be
this subspecies.

Paoliella chiangae van Harten and Ilharco Only alatae are described; they
have head, thorax and legs brown-black, abdomen greyish-brown with dark
tubercles, BL 1.2-1.6 mm (van Harten and Ilharco, 1972). Living singly or in
small colonies on undersides of young leaves of Combretum sp. (?psidioides)
in Angola.

Paoliella commiphorae (Doncaster) All viviparae alate, yellowish, thickly

dusted with grey wax; BL c. 1.5 mm. On Commiphora zimmermannii in Kenya
and Malawi. Similar aphids collected on C. fischeri in Tanzania were described
as a subspecies, C. commiphorae persimilis (Eastop, 1955b). Alate males of
this latter form have been trapped in January-March (BMNH colln).
 Paoliella 795

Paoliella delottoi (Hille Ris Lambers) Apterae brownish, BL 0.9-1.2 mm,

feeding on undersides of shoot tips of Commiphora africana in Eritrea
and Nigeria. Life cycle unknown. Aphids collected on C. dulcis in Namibia
(BMNH colln, leg. H.D. Brown) are the same or a closely related species.

Paoliella echinata Eastop Apterae black with white dorsal processes; BL

c. 0.8 mm. Immatures yellow. On Commiphora pilosa in Tanzania and South
Africa. Biology unknown.

Paoliella harteni Ilharco Only alatae known, orange-reddish appearing rose-

coloured due to fine pulverulence; BL 1.5-1.9 mm. On both sides of leaves
of Terminalia spp. (brachystemma, sericea) in Angola (Ilharco, 1971). Van
Harten and Ilharco (1972) described the alate male and apterous ovipara,
collected in July. Viggiani (1987) described a new aphelinid parasitoid from
this aphid.

Paoliella hystrix Theobald, Plate 7e Colour of aptera in life not recorded;

BL 0.5-0.8 mm. Alatae and life cycle unknown; only known from original
collection on Commiphora savojae in Somalia (Theobald, 1928).

Paoliella longirostris Quednau Only alatae known, colour in life unrecorded,

BL c. 1.5 mm. Described from an alata trapped in Zimbabwe (Quednau, 1974)
and subsequently collected in Burundi on Combretum sp. (Remaudière and
Autrique, 1985).

Paoliella nachensis (Eastop) All viviparae alate, head and thorax dark,
abdomen yellow with dark dorsal processes; BL 1.1-1.4 mm. Living singly
on undersides of leaves of Combretum spp. (incl. apiculatum, ?psidioides).
Recorded from Angola, Burundi and Tanzania. Life cycle unknown.

Paoliella namaachae Ilharco Only alatae known, with pale yellow-orange

abdomen and slightly darker head and thorax; BL 0.9-1.2 mm. Living
solitarily on undersides of leaves of Combretum molle in Mozambique.
Ilharco (1976) discussed differences from the closely-related P. nachensis.
Life cycle unknown.

Paoliella nirmalae (David) Appearance in life unknown, probably rather

pale; BL of aptera 1.1-1.4 mm. On Terminalia arjuna in Mysore, India
(David, 1969). Life cycle unknown.

Paoliella papillata (Hall) Plate 7f All viviparae alate, very dark brown; BL
1.4-1.5 mm. Immatures pale reddish-brown with white dorsal processes (Hall,
1932). On Combretum spp. (incl. apiculatum, molle), widely distributed in
Africa south of the Sahara. Doncaster (1954) described the alate male and
apterous oviparae from amongst the type material, collected in Zimbabwe in
December.
796 Paracletus

Paoliella terminaliae (Hall) All viviparae alate, brick red to very dark but
with tip of abdomen always pale; BL c. 1.5 mm (Hall, 1932). On Terminalia
sericea in southern Africa (Angola, Malawi, South Africa, Zimbabwe). Life
cycle unknown. Eastop (1956) described kenyensis as a subspecies, from a
single alata trapped in Kenya.

Paoliella ufuasi (Eastop) Only alatae known, appearance in life not

described; BL 1.2-1.4 mm. Described from trapped alatae in Tanzania
(Eastop, 1955b), but specimens collected on Commiphora pilosa in Kenya
are believed to be this species (Quednau, 1974), and alatae collected on
C. angolensis in South Africa are very similar (BMNH colln, leg. A.L.
Capener). Life cycle unknown.

Paoliella wettsteini Quednau Apterae pale yellow with white dorsal pro-
cesses; BL 1.0-1.2 mm. Alatae very pale with head and thorax only slightly
dusky. On leaves of Combretum gueinzii (= C. molle) in South Africa
(Quednau, 1964). Life cycle unknown.

PARACLETUS van Heyden Pemphiginae: Fordini

About 4 species resembling Forda except for the longer outer margin of the
hind coxae. Only P. cimiciformis is known to complete the holocycle with
galls on Pistacia; sexuparae of P. donisthorpei have been recorded from
P. terebinthus (Roberti, 1939), but the gall-living generations of this species
are unknown.

Paracletus cimiciformis von Heyden The galls on Pistacia (khinjuk, palaes-

tina, terebinthus) are flat folds of the edges of the leaves, retaining the
original leaf colour. The fundatrix - globose, yellow, with 6-segmented anten-
nae, BL c. 1.5mm - has never been properly described. Second generation
are all alate, BL 1.7-2.4 mm, emerging from the gall in September-October
to found colonies on roots of various Gramineae. Sexuparae return to
Pistacia in April-June; these and the sexual morphs were fully described by
Roberti (1939). Recorded from Pistacia in southern France, Italy, Iran,
Israel and Turkey. Anholocyclic populations on roots of grasses occur
outside the range of the primary hosts (see B & E, 1984, p. 323). Predation
by birds on gall populations was observed by Burstein and Wool (1992).
2n = 16.
 Paracolopha 797

PARACOLOPHA Hille Ris Lambers Pemphiginae: Eriosomatini

Two species in East Asia closely related to Colopha, but the media of the
forewing of alatae from galls is unbranched and the apterae on the secondary
hosts (bamboos) always have siphunculi. The life cycle of only one species is
known. Akimoto (1985a) gave a full account.

Paracolopha morrisoni (Baker) Gall on Zelkova serrata is a club-shaped

outgrowth from the upper surface of the leaf lamina between veins, with
a narrow stem and a swollen, roughened apex (Fig. 125C, p. 655). Alatae
emerge in June-July and colonize roots of bamboos (Sasa spp.). Sexuparae
return to Zelkova in September-October. Moritsu's (1958) record from
Celtis sinensis var. japonica (as Tetraneura nishiyae) was probably based on
vagrant sexuparae. Galls on Zelkova occur in Japan and southern China (as
Tetraneura zelkovisucta; Zhang, 1980b), and presumably anholocyclic
populations have been introduced into USA, where it is recorded from the
roots of various bamboos (Phyllostachys, Arundinaria) in Maryland, South
Carolina and southern California (Hille Ris Lambers, 1966d). This species has
a complex synonymy; commonly used names in the past have been Colopha
moriokaensis (Monzen) and Dryopeia nishyae (Matsumura, 1931, nec. 1917).
2n = 10*.

PARADOXAPHIS Sunde Aphidinae: Aphidini (?)

One species in New Zealand, resembling Aphis but lacking lateral tubercles
on ABD TERG 7, and often also lacking them on ABD TERG 1.

Paradoxaphis aristoteliae Sunde Apterae reddish-brown with black head

and dusky thorax; BL 1.5-2.0 mm. On leaves of Aristotelia serrata in New
Zealand (Sunde, 1987). Biology unknown. 2n = 8*.

PARANIPPONAPHIS Takahashi
Hormaphidinae: Nipponaphidini

One species on Quercus in Japan, close to Nipponaphis but without siphun-

cular pores.
798 Paraprociphilus

Paranipponaphis takaoensis Takahashi Apterae nearly circular, with sides

of body perpendicular ('pill box-shaped'), blackish-brown, BL 1.5-2.0 mm.
Other morphs unknown. On branches of an unidentified Quercus sp. in
Japan (Takahashi, 1959b). Other morphs and life cycle unknown.

PARAPROCIPHILUS Mordvilko Pemphiginae: Pemphigini

Three species, two nearctic and one palaearctic. The nearctic species have
host alternation between Acer and Alnus; the other lives on Alnus and pro-
duces sexuparae in autumn, but its primary host generations are unknown.
Remaudière and Muñoz Viveros (1985a) summarized differences between the
species.

Paraprociphilus baicalensis (Cholodkovsky) Apterae brownish, secreting

copious white wax wool; BL c. 3.1 mm. On undersides of leaves and shoots of
Alnus spp., across Asia from the Urals to Japan. Apparently entirely anholo-
cyclic on Alnus, although sexuparae are produced in autumn (Mordvilko,
1935). 2n = 12*.

Paraprociphilus mexicanus Remaudière and Muñoz Viveros Apterae dark-

coloured, entirely covered with a dense waxy secretion, with long wax
filaments projecting posteriorly; BL 2.9-3.6 mm. On undersides of leaves
of Alnus firmifolia in Mexico (Remaudière and Muñoz Viveros, 1985a).
Sexuparae are produced in autumn, and there is at least a partial migration
to Acer negundo, from which fundatrices and emigrant alatae were recently
described (Remaudière and Muñoz Viveros, 1993).

Paraprociphilus tessellatus (Fitch) Brownish aphids living beneath a copious

secretion of white woolly wax. Fundatrices and their alate progeny develop
on undersides of leaves of Acer saccharinum in May-July, causing slight leaf
curl. Migration occurs to Alnus, where colonies (BL of aptera 3.6-4.4mm)
are formed along the branches. Alate sexuparae return to Acer in September,
but anholocyclic overwintering also occurs under fallen leaves or in loose earth
at the base of alder trees (Patch, 1911; Mordvilko, 1935). Widely distributed
in North America. A neuropteran predator (Chrysopa slossonae) lays its eggs
specifically in relation to colonies of P. tessellatus (Bristow et al., 1988).
2n = 6*.
 Parathoracaphis 799

PARATHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini

Five species in East and Southeast Asia differing from Thoracaphis in the
complete fusion of prosoma and abdominal plate, different cuticular orna-
mentation, spine-like marginal hairs and absence of siphunculi. Four species
are on Fagaceae; the host of the fifth is only tentatively recorded as Myrica.
Only apterous viviparae are known. Some of the species may have unrecog-
nized gall-forming generations on Distylium.

Parathoracaphis cheni (Takahashi) Apterae black, oval, flattened, strongly

sclerotized, without wax; BL c. 0.9mm. Sedentary on undersides of leaves of
?Myrica rubra (host is queried in original description) in China (Chekiang).
Life cycle unknown.

Parathoracaphis elongata (Takahashi) Apterae black, elongate oval, strongly

sclerotized, with numerous wax filaments dorsally; BL c. 0.5 mm. On under-
sides of leaves of Quercus sp. and other unspecified Fagaceae, in Thailand
(Takahashi, 1941). Life cycle unknown.

Parathoracaphis gooti (Takahashi) Apterae black, broadly oval, flattened,

sclerotized, with much white marginal wax; BL c. 1 mm. On undersides of
leaves of Quercus sp. in Malaya (Takahashi, 1950). Life cycle unknown.

Parathoracaphis kayashimai (Takahashi) Apterae presumably black, elon-

gate oval, strongly sclerotized, perhaps without wax (not stated); BL c. 0.5 mm.
On undersides of leaves of Quercus sp. in Malaya (Takahashi, 1950). Life
cycle unknown.

Parathoracaphis setigera (Takahashi) Apterae black, elongate oval, strongly

sclerotized, without wax; BL 0.5-0.7 mm. In loose clusters on undersides of
leaves of Quercus spp. (glauca, myrsinaefolia) and Lithocarpus sp. in Japan,
China (Sikong) and Taiwan (Tao, 1966). In Japan apterae occur on Quercus
throughout the year (Takahashi, 1958a).

PARATHORACAPHISELLA Pramanick, Samanta and

Raychaudhuri Hormaphidinae: Nipponaphidini

One Indian species on Quercus, apparently close to Metathoracaphis. Only

apterous viviparae are known.
800 Patchiella

Parathoracaphisella indica Pramanick, Samanta and Raychaudhuri Apterae

dark brown, with margin darker, oval; BL c. I.i mm. On Quercus sp. at
Manipur, India (Pramanick et al., 1983).

PATCHIA Baker Drepanosiphinae: Phyllaphidini

Two North American species on Fagaceae, perhaps closely related to Lachno-

chaitophorus, although the apterae (at least in the type species of the genus)
have separate head and prothorax.

Patchia virginiana Baker Apterae broadly oval, rather uniformly velvety-

black, secreting wax; BL 1.8-2.3 mm. Alatae are brown with a large black
dorsal abdominal patch and thickly-bordered forewing veins. On bark of
Castanea spp. in USA (Florida, North Carolina, Pennsylvania, Virginia).
Apparently monoecious holocyclic; oviparae were collected in September
(BMNH colln, leg. J.O. Pepper).

Patchia winforii Miller Apterae dark brown to black; BL c. 1.7 mm. Alatae
have black dorsal abdominal cross-bands and thickly-bordered wing veins.
Forming dense clusters on twigs and leaf petioles of Quercus palustris in
Massachusetts, USA (Miller, 1933b). Life cycle unknown. Not recorded since
the original description.

PATCHIELLA Tullgren Pemphiginae: Pemphigini

One palaearctic species host-alternating between Tilia and Araceae (B & E,

1984, p. 323).

Patchiella reaumuri (Kaltenbach) Forming large leaf-nest galls on Tilia spp.

in spring by twisting and stunting the terminal growth and clumping the leaves.
Fundatrices are plump-bodied, yellowish-brown to greenish; BL c. 3.5mm.
All second generation are alate, with abdomen green to yellow-green, BL
2.1-2.7 mm, migrating to found colonies on tubers of Arum spp. (Roberti,
1939). Recorded from Tilia in England, Italy, Bulgaria, Spain and Turkey,
and anholocyclic populations occur on roots of Araceae elsewhere (e.g.
Hawaii, Solomon Islands). Pashchenko (1984) described as a subspecies,
P. reaumuri orientalis, from Tilia amurensis in Siberia; this is presumably
the insect referred to as orientalis Mordvilko in Shaposhnikov (1955).
 Pemphigus 801

PAULIANAPHIS Essig Greenideinae: Cervaphidini

One species on Madagascar related to Eonaphis and Schoutedenia, but with

two pairs of dorsal processes on ABD TERG 7 and a different host associa-
tion. Remaudière (1988) gave a full account.

Paulianaphis madagascariensis Essig Apterae broadly oval (almost glo-

bular), colour in life not recorded; BL 1.7-2.1mm. In galls on leaves of
Apocynum [label on type slide in BMNH colln, although host given as
'Apocynaceae' in Essig (1958); host plant and locality data given in error
under Eonaphis pauliani (see Hille Ris Lambers, 1966b)]. Alatae have
thickly-bordered forewing veins. Only known from Madagascar. Life cycle
unknown.

PEMPHIGUS Hartig Pemphiginae: Pemphigini

Over 70 species have been described, of which 46 are known to form galls on
leaves or twigs of Populus (subgenus Eupopulus). Alatae emerging from these
galls have a black head and pterothorax and a rather elongate, yellow-green
to grey-green, wax-dusted, abdomen. The life cycles of 17 of these gall-
formers are known; four are monoecious on Populus, the alate sexuparae
being produced in the galls, and in the other 13 species the alate progeny of
the fundatrix migrate to various herbaceous secondary hosts (in Cruciferae,
Compositae, Chenopodiaceae, Euphorbiaceae, Ranunculaceae and Umbelli-
ferae), where they found colonies on the roots, or occasionally in woolly wax
masses above soil level. Most of the species known only from their root-
feeding generations feed on herbs in the above-named families, but at least
one species feeds on Salix roots. The species are distributed rather evenly
throughout the northern hemisphere (see B & E, 1984, p. 324). Accounts are
available for Britain (Furk and Prior, 1975); European gall-makers on
Populus nigra (Lampel, 1960); Fennoscandia and Denmark (Heie, 1980);
India (Ghosh, 1984b); eastern Russia (Pashchenko, 1988b); Japan (Aoki,
1975); China (Zhang and Zhong, 1979, 1983b); and North America (Palmer,
1952; Harper, 1959a, b; Lange, 1965; Smith, 1985).

Pemphigus betae Doane (= balsamiferae Williams) Galls are yellowish-

green sometimes tinged with red, pocket-like, formed usually on the underside
of the poplar leaf near its base, on or adjacent to the mid-rib, with little or
no deformation of the leaf (Harper, 1959a; Whitham, 1978). The opening can
be either on the upper side of the leaf or underneath along the mid-rib.
802 Pemphigus
 Pemphigus 803

Primary hosts are Populus angustifolia, P. balsamifera, or occasionally

P. deltoides var. occidentalis or P. trichocarpa. Alatae (BL 1.5-2.1mm)
leave galls in late June-early August (Harper, 1959b), to colonize roots of
Chenopodiaceae (B & E, 1984, p. 324). Anholocyclic populations also persist
all year on secondary hosts (see Moran and Whitham, 1988; Moran, 1991).
Widely distributed in North America. The life cycle was studied by Harper
(1963) in southern Alberta, and by Moran and Whitham (1988) in Utah. This
aphid's evolutionary ecology has been studied in some detail, including the-
territorial behaviour of first instar fundatrices in selection of galling sites
(Whitham, 1979), population dynamics in relation to galling site (Whitham,
1978, 1980), genetics of resistance in the host plant (Paige and Capman, 1993)
and discrimination between host trees by returning sexuparae in autumn
(Moran and Whitham, 1990).

Pemphigus borealis Tullgren Galls on Populus laurifolia are green, ellip-

tical, hazelnut-sized, usually in groups of 2-10 on stem at bases of twigs or
leaf petioles (Fig. 131 K). Other recorded primary hosts are P. balsamifera
(planted), P. koreana, P. maximowiczii and P. suaveolens. Fundatrix is grey-
green with slight yellowish tinge. Alatae (BL 1.5-2.5 mm) emerge in June-
August and migrate to colonize roots of Bidens spp. (Dolgova, 1970). Return
migration occurs in August-September. Northern palaearctic, from Sweden
to China and Japan. In East Asia there is probable confusion with P. dorocola
(q.v.), which is described from very similar galls on P. maximowiczii and
is possibly a synonym (Aoki, 1975). Pemphigus yangcola, described from
Populus sp. in China (Chang and Zhong, 1979c), is also very similar and a
possible synonym. 2n = 20*.

Pemphigus bursarius (Linnaeus) Galls on Populus spp. (most commonly

P. nigra) are yellowish or reddish when mature, purse-shaped, always formed
on leaf petioles (Fig. 131L). Fundatrix is greyish-green, slightly wax-dusted.
Alatae (BL 1.6-2.5 mm) leave the gall through a lateral opening over an

Fig. 131. Galls on Populus. A, Epipemphigus sanpopuli on P. cathayanae (after Zhang and Zhong,
1983b); B, E. imaicus on P. ciliata (after Habib and Ghani, 1970); C, Gootiella tremulae on P. tremula
(after Danielsson, 1976); D, Mordwilkoja vagabunda on P. deltoides (after Britton, 1923); E, Pachypappa
marsupialis on P. nigra (after Börner and Heinze, 1957); F, Pachypappa populi on P. tremula (after
Danielsson, 1976); G, Pachypappa tremulae on P. tremula (after Tullgren, 1909); H, Pachypappella
lactea on P. tremula, with colony overflowing onto outside of gall (after Danielsson, 1976); I, Thecabius
affinis (fundatrix gall) on P. nigra (after Börner and Heinze, 1957); J, Th. lysimachiae on P. nigra (after
Börner and Heinze, 1957); K, Pemphigus borealis on P. laurifolia (after Tullgren, 1909); L, Pemphigus
bursarius on P. nigra (after Börner and Heinze, 1957); M, Pemphigus immunis on Populus sp. (after
Theobald, 1915); N, Pemphigus monophagus on P. angustifolia (after Aoki, 1975); 0, Pemphigus passeki
on P. nigra (after Prinsen, 1990); P, Pemphigus populinigrae on P. nigra (after Börner and Heinze,
1957); Q, Pemphigus populitransversus on P. deltoides (after Palmer, 1952); R, Pemphigus protospirae
on P. nigra (after Danielsson, 1976); S, Pemphigus spyrothecae on P. nigra (after Danielsson, 1976);
T, Pemphigus vesicarius on P. nigra (after Lichtenstein, 1855).
804 Pemphigus

extended period from late May to September, with peak emergence in late
June-July (northern hemisphere). They found colonies mainly on roots of
Compositae (except for populations in Australia and New Zealand which are
mainly recorded from Chenopodiaceae and Polygonaceae, and are perhaps a
separate species). Return migration of sexuparae to poplar is in September-
October, but anholocyclic overwintering on roots of secondary host is com-
mon. In Europe, northwest, southwest and Central Asia, Australia and
New Zealand, North and southern Africa, North and South America. Dunn
(1960a) studied gall formation and (1960b) reviewed natural enemies. Alleyne
and Morrison (1978a, b) gave accounts of the bology and natural enemies in
Quebec. See also B & E, 1984, p. 325. 2n = 20.

Pemphigus chomoensis Chang Described from fundatrices and immatures

only, in a mid-rib gall similar to that of Epipemphigus niisimae, on Populus
sp. in Tibet (Chang and Zhong, 1979c). Possibly this is a synonym of
Epipemphigus imaicus.

Pemphigus circellatus Zhang and Zhong Large galls are formed on the
branches of Populus tremula var. davidiana in China (Yunnan). Emigrant
alatae (BL c. 1.8mm) were collected in May (Zhang and Zhong, 1985a).
Life cycle unknown.
Pemphigus cylindricus Zhang Only fundatrices are described, from a gall
on an unidentified Populus sp. in Tibet (Zhang and Zhong, 1981b).
Pemphigus dorocola Matsumura The galls on twigs of Populus maximo-
wiczii in Japan are pouch-shaped, reddish when mature, with a transverse
apical slit (Aoki, 1975). Also recorded from P. koreana and P. suaveolens.
Fundatrix green; alatae (BL 1.9-2.3 mm) emerge over an extended period
(late July-November). Secondary host unknown. In Japan, Korea and Siberia
(Pashchenko, 1988b). Aoki (1978, 1980a) described aggressive and gall-
cleaning behaviour by first instar larvae, and Aoki and Kurosu (1991d)
described an aptera producing abnormal progeny in the gall. [P. dorocola is
separated in the key to gall-formers on Populus according to the description
by Aoki (1975), who does not recognize the presence of P. borealis in Japan.
Alatae in the BMNH colln from galls on Populus maximowiczii in Japan and
Korea all have antennal sensoriation like that of P. borealis, and further work
may show P. dorocola to be a synonym of P. borealis, as suggested by
Takahashi (1920).] 2n = 20 (Blackman, 1986).
Pemphigus gairi Stroyan One of three palaearctic species producing similar
pouch-shaped galls on or near the mid-rib on the upper sides of leaves of
Populus nigra (incl. var. italica), with an opening to the underside of the leaf
(the other two species are P. phenax and P. populinigrae). Fundatrix dull pale
green. Alatae (BL 1.5-2.2 mm) emerge in June-July and migrate to colonize
roots of Aethusa cynapium (Stroyan, 1964b). Sexuparae return in October.
Only recognized from England, but probably more widely distributed in
Europe.
 Pemphigus 805

Pemphigus ignotus Habib and Ghani Recorded as forming leaf galls on

Populus ciliata in Pakistan (Habib and Ghani, 1970), but no description is
given and this is therefore a nomen nudum.

Pemphigus immunis Buckton (= lichtensteini Tullgren) The galls are quite

large (up to 4cm max. diam.), walnut-like in shape and surface texture, thick-
walled, with an apical opening, on twigs or branches of Populus nigra,
P. euphratica, and less commonly P. ciliata (Fig. 131M). Fundatrix is glaucous
green (Roberti, 1938). Alatae (BL 2.2-2.5 mm) emerge from late April to early
August and found colonies on roots of annual Euphorbia spp. (helioscopia,
peplus; Hille Ris Lambers, 1973b). In Europe, North Africa, southwest and
Central Asia, Pakistan, northwest India, and China (Tseng and Tao, 1936;
as P. napaeus). Habib and Ghani (1970) studied natural enemies in Pakistan
and Bhagat (1982a) reported a parasitoid and hyperparasitoid in Kashmir.
2n = 10 according to Pal and Khuda-Bukhsh (1982).

Pemphigus iskanderkuli Narzikulov Galls on Populus dentata when mature

are pinkish to red, rounded, smooth, walnut-sized, with apical opening,
situated on upper leaf surface at junction of petiole and lamina (Narzikulov,
1957). BL of alata 2.7-3.5 mm. In Tadzhikistan. Life cycle unknown. The
alatae seem indistinguishable from those of P. napaeus, and Doncaster (1969)
suggested that the two might be conspecific, but for reported differences
in the morphology of the fundatrices, and position and colour of the galls.
Populations in Kirgizia on P. laurifolia described by Gabrid (1989) as
Pemphigus sp. aff. iskanderkuli would appear to be large specimens of
iskanderkuli, although the galls illustrated by Gabrid appear very irregular
in shape.

Pemphigus junctisensoriatus Maxson Galls on Populus deltoides are pale

yellowish-green, composed mostly of swollen leaf petiole around broadened
base of leaf lamina, with exit hole in the form of a long slit between petiole
and leaf (Maxson, 1934). Fundatrix is pale yellowish-white, with brownish
head and tibiae. Alatae (BL 1.4-1.8 mm) emerge in June-July. Heteroecious;
secondary host unknown. In USA and eastern Canada.

Pemphigus knowltoni Stroyan Galls are green, pouch-like, of variable

shape, on undersides of leaves of Populus angustifolia alongside the mid-rib
at the base of the lamina (Stroyan, 1970). Fundatrix and immatures in gall
are blackish-green. Alatae (BL 1.8-2.1 mm) are mature in the gall in July.
Described from Utah, USA. Heteroecious; secondary host unknown.

Pemphigus laurifoliae Dolgova Galls occur on Populus laurifolia in the

Altai region of Russia. Alatae (BL c. 2.1 mm) emerge in July (Dolgova, 1973).
[Original description not available; keyed on the basis of alate specimens
(paratypes?) collected by L.P. Dologova on l.vii.71 (BMNH colln, leg.
DHRL). Pemphigus laurifoliae Holman and Szelegiewicz, 1972 (nomen
nudum) is P. mongolicus Holman and Szelegiewicz, 1974a.]
806 Pemphigus

Pemphigus longicornus Maxson Galls elongate, thin-walled, on upper sides

of leaves of Populus deltoides. Fundatrix pale yellowish-white. Alatae (BL
1.7-2.1mm) emerge in May-June. In eastern and southern USA. Hetero-
ecious, secondary host unknown (Maxson, 1923). [According to Smith (1985),
there is also an undescribed Pemphigus sp. on P. deltoides in North Carolina
that causes elongate to slightly globular galls on the upper sides of the leaves,
and has alatae with antennae intermediate in length and antennal sensoriation
between longicornus and populivenae.]
Pemphigus mangkamensis Zhang Described from apterous viviparae from
an unidentified Populus sp. in Tibet (Zhang and Zhong, 1981b) and said to
be similar to P. chomoensis. If the specimens are not fundatrices then this
would indicate absence of host alternation.
Pemphigus matsumurai Monzen Gall is globular, at base of mid-rib on
upper side of leaf, concolorous with mid-rib or slightly reddish (Aoki, 1975;
Zhang and Zhong, 1983b). Recorded from Populus ciliata, P. koreana,
P. maximowiczii and P. suavolens. Fundatrix is green. Alatae (BL 1.7-
2.7mm) migrate in June-July to colonize roots of Thalictrum spp. In East
Asia (northern India, Pakistan, Mongolia, Siberia, Korea, China, Japan).
Also in Tadzhikistan, as Pemphigus montanus, described from Populus
tadzhikistanica (Narzikulov, 1957), is clearly a synonym. Aoki and Kurosu
(1989) studied the influence of parasitization of the fundatrix on development
of the gall and subsequent generations. Bhagat's (1982a) record of a parasitoid
of P. immunis possibly should be referred to matsumurai, as this is the only
species known to form leaf galls on P. ciliata. 2n = 12 (an unusual chromo-
some number for a Pemphigus; Blackman, 1986).
Pemphigus microsetosus Aoki A large closed pouch-gall is formed as an
outgrowth of the leaf of Populus maximowiczii, taking in the whole leaf
when mature; lignified galls remain on the tree after leaf-fall (Aoki, 1975).
Fundatrix yellowish-green. Alatae (BL 2.1-2.5 mm) are adult from late July
and leave gall through several secondary exit holes. In Japan and eastern
Siberia (Pashchenko, 1988b). Heteroecious; secondary host unknown. 2n =
22 (Blackman, 1986).

Pemphigus mongolicus Holman and Szelegiewicz Gall is globular, pouch-

like, on upper side of leaf of Populus laurifolia near the base of the mid-rib;
green, becoming tinged with red when mature (Holman and Szelegiewicz,
1974a). Fundatrix is palish green, producing a generation of dark green
apterae in the galls. Monoecious; alate sexuparae (BL 1.6-2.0 mm) leaving
galls in early August give birth to sexuales in bark crevices on the trunk. In
Mongolia. Possibly 'P. semenovi', noted by Mordvilko (1935) as monoecious
on P. maximowiczii and P. suaveolens in Siberia, but apparently a nomen
nudum, is this species.

Pemphigus monophagus Maxson Gall is globular, pale greenish-yellow,

consisting of a swelling of upper side of base of leaf lamina with petiole
 Pemphigus 807

broadened and somewhat twisted, and a slit-shaped opening underneath

between petiole and leaf (Fig. 131N; and Maxson, 1934). Fundatrix honey-
yellow to pale olive green, producing a generation of creamy-white apterae in
the gall. Monoecious, alatae (BL 1.4-1.9 mm) emerging from galls in August-
September to deposit sexuales on bark of trunk. On Populus angustifolia,
P. balsamifera and P. trichocarpa in western North America. Aoki and
Kurosu (1988) discussed the evolution of monoecy in this species.

Pemphigus montanus Narzikulov See P. matsumurai.

Pemphigus mordvilkoi Cholodkovsky Galls, formed on the twigs of Populus

ciliata, are smooth, globular, sessile, yellowish-green to dark greenish-brown
(Chakrabarti et al., 1985). Fundatrix pale green. The life cycle is unusual and
requires further study. In Pakistan, alatae emerge from the galls over an
extended period from August until January of the next year (BMNH colln).
According to A.K. Ghosh et al. (1981), the alatae emerging from galls just
before onset of winter in northwest India hibernate in bark crevices of the
same tree, or in other sheltered places; however, these alatae are not sex-
uparae, so the monoecious life cycle suggested by them seems unlikely.
Apterous exules and sexuparae likely to be those of P. mordvilkoi have been
collected in Punjab in April on roots of Acroclinum roseum (BMNH colln,
leg. A.S. Sohi); similar sexuparae have also been collected in other parts of
northern India and Nepal in March-April (BMNH colln), and sexuparae and
sexuales were observed in Pakistan in April-May (Habib and Ghani, 1970).
The oviparae laid eggs which hatched in six-nine days, and the fundatrices
formed galls in May-June. Thus a two-year life cycle with alternation between
P. ciliata and Acroclinum is indicated. 2n = 20*.

Pemphigus nainitalensis Cholodkovsky Galls about the size and shape of a

hazelnut, sessile, on twigs of Populus ciliata, which are thickened in the region
of the gall (Cholodkovsky, 1912). Only known from the original descriptions
of fundatrix and alata, from galls collected in Uttar Pradesh, India.

Pemphigus napaeus Buckton Galls on twigs of an unidentified Populus sp.

were smooth, roundish or of irregular shape, shining green 'slightly veined
with brown, and variegated with yellowish patches' (Buckton, 1896); but
see comments by Doncaster (1969, p. 162). In Kashmir, India. There are
no other authenticated records and contrary to some authors (e.g. A.K.
Ghosh et al., 1981), the host species is unidentified. Reports of napaeus in
Taiwan (e.g. Tseng and Tao, 1936) should probably be referred to immunis.
Doncaster (1969) remounted and redescribed the type material. See also
under P. iskanderkuli.

Pemphigus nortonii Maxson Galls on Populus deltoides, especially var.

occidentalis (= sargentii), are pale yellowish-green, formed by a swelling
at the base of the leaf lamina, twisted on the flattened, broadened petiole;
resembling those of P. populicaulis (Palmer, 1952), but with a long slit-like
808 Pemphigus

opening (Harper, 1959a). Alatae emerge over an extended period (June-

September; Harper, 1959b; Grigarick and Lange, 1968). Heteroecious, but
secondary host unknown. Apparently widely distributed in USA, and in
Alberta, Canada. Apart from the greater number of rhinaria on ANT V and
VI this species closely resembles P. junctisensoriatus. Populations from
Alberta (Harper, 1959a, b) seem to be intermediate in this respect and the
distinction between these two species needs to be verified.

Pemphigus passeki Börner Galls are formed on leaves of Populus nigra

(incl. var. italica) by swelling of the basal half of the mid-rib, the swelling
being broadest towards the junction with the petiole (Fig. 131O; and Prinsen,
1990). Alatae (BL c. 2.1 mm) fly in June-August to colonize roots of caraway
(Carum carvi). Northern palaearctic; northwest and Central Europe, east to
Altai region of Russia (BMNH colln, leg. L.P. Dolgova). Spring migration
was studied by Prinsen (1990). 2n = 22 (Gut, 1976).

Pemphigus phenax Börner and Blunck One of a complex of species in

Europe producing similar mid-rib galls on upper sides of leaves of Populus
nigra (incl. var. italica), with a slit-like opening underneath the leaf. The galls
of P. phenax are reddish, tend to be larger than those of P. gairi, and more
elongate (spindle-shaped) than those of P. populinigrae, somewhat more
wrinkled and often tinged with yellow laterally (Stroyan, 1964b). Fundatrix
is dull pale green when adult. Alatae (BL 1.5-2.4 mm) fly in June-August
to found colonies on roots of carrot (Daucus carota). In Europe (England,
Denmark, France, Germany, The Netherlands, Sweden). Populations also
overwinter anholocyclically on carrot roots.

Pemphigus plicatus Dolgova Galls on Populus laurifolia in the Altai region

of Russia. Alatae (BL c. 1.8mm) collected in July. Original description not
available. [Keyed on basis of specimens collected by Dolgova on l.vii.71
(BMNH colln, leg. L.P. Dolgova.]

Pemphigus populi Courchet Galls on Populus nigra (incl. var. italica) are
globular outgrowths of the mid-rib, not larger than a hazelnut, becoming club-
shaped with the basal part narrower than the apex when mature, situated near
the base on the upper side of the leaf. Alatae (BL 1.4-2.5 mm) emerge in
June-July through a secondary opening at the top of the gall, migrating to
found colonies on roots of Melilotus altissima and possibly other Leguminosae
(Furk and Prior, 1975). Sexuparae return to P. nigra in September-October.
In Europe, southwest and Central Asia, and east to China (Chang and Zhong,
1979c). Furk and Prior (1975) reported parasitization of fundatrices by
Monoctonia pistaciaecola.

Pemphigus populicaulis Fitch Gall is yellowish-green to creamy, formed by

swelling of base of leaf lamina, twisted around broadened petiole, rather like
that of P. nortonii and P. populiglobuli but with a small, rounded exit hole
on underside (Palmer, 1952; Harper, 1959a). Especially associated with
 Pemphigus 809

P. deltoides and P. deltoides var. occidentalis (= sargentii), but also recorded

from other cottonwoods and P. tremuloides. Fundatrix pale yellowish to
whitish-green. Alatae (BL 1.7-2.0 mm) emerge in July-September. The secon-
dary host has not definitely been established; Glendenning (1924) reported that
it was Oenanthe sarmentosa, but did not provide evidence for this assertion.
Hottes and Frison (1931) suggested that P. brevicornis Hart, described from
corn roots, was the sexupara of populicaulis. Widely distributed in North
America. Setzer (1985) studied gall mortality. Sokal et al. (1980) and Bird
et al. (1981) studied geographical variation in eastern North America using
multivariate techniques.

Pemphigus populiglobuli Fitch Gall is green, often tinged with red and/or
yellow when mature; globular, conical or bullet-shaped, formed by swelling
of base of leaf lamina and thickening of petiole, the exit hole being a slit
on the underside between leaf and petiole (Maxson and Knowlton, 1929).
Primary hosts are Populus angustifolia and P. balsamifera, more rarely
P. trichocarpa. Fundatrix is variously reported to be dark green (Palmer,
1952) or pale yellowish-white (Maxson, 1934). Alatae (BL 1.5-2.3 mm) leave
gall in June-early July (Alberta; Harper, 1959b) for an unknown secondary
host. Widely distributed in North America. Differences from P. populicaulis
and other closely-related species are discussed by Maxson and Knowlton
(1929).

Pemphigus populinigrae (Schrank) (=filaginis Boyer de Fonscolombe) Galls

on Populus nigra are pouch-shaped (Fig. 131P), similarly situated to those of
P. phenax and P. gairi, on upperside of leaf lamina alongside mid-rib with
opening along original line of invagination on underside of leaf. Gall forma-
tion was described by Dunn (1960a). Mature galls are rounded and smooth
on top and of a dull reddish colour without much tinge of yellow (Stroyan,
1964b). Fundatrix is dark green. Alatae (BL 1.8-2.9 mm) emerge in June-
August and migrate to found colonies above ground on Filago and Gnapha-
lium, often in the tomentum of the inflorescences. Sexuparae return to
P. nigra in September-October, but anholocyclic overwintering apparently
occurs where the primary host is not available (Mordvilko, 1935). Throughout
Europe and in North Africa, southwest Asia, Central Asia and across to
eastern Siberia (Pashchenko, 1988b). Records from China (Tao, 1958) may
be due to confusion with Epipemphigus niisimae. 2n = 22.

Pemphigus populiramulorum Riley Galls are formed by swelling of new

growth of twigs, or rarely at the bases of petioles; they are quite large (up to
2.5 cm diam.), rounded, green to brownish, with a transverse or vertical slit-
shaped opening which often has protruding lips. Primary host is usually
Populus deltoides var. occidentalis (= sargentii); also recorded from P.
balsamifera and P. fremontii. Fundatrix is pale creamy-yellow (Palmer, 1952).
Alatae (BL 2.2-2.5 mm) leave galls in July-September (Harper, 1959b), flying
to an unknown secondary host. In western North America (Utah, Colorado,
South Dakota).
810 Pemphigus

Pemphigus populitransversus Riley Plate 6c Gall is formed by swelling

and bending of leaf petiole of Populus deltoides (incl. var. occidentalis =
sargentii); it is green, globular to ovoid, with a usually transverse slit (Fig.
131Q). Sometimes recorded from other Populus spp. (acuminata, angusti-
folia, balsamifera, fremontii, nigra, tremuloides, trichocarpa). However,
some of these records, particularly those from western cottonwoods, may
apply to other species; e.g. Smith (1985) noted that the galls commonly found
on P. fremontii in California were not of the form typical of eastern populi-
transversus. Fundatrix is whitish to pale dirty greenish-yellow (Palmer, 1952).
Alatae (BL 1.7-2.0 mm) emerge in July-September and found colonies on
roots of Cruciferae (Jones and Gillette, 1918), where it can be a serious pest.
[But not in California, suggesting again that the populitransversus-like alatae
produced in galls on P. fremontii are a distinct species, perhaps with Rumex
as secondary host (D. Hille Ris Lambers, pers. comm.).] Sexuparae return
to cottonwood in September-October to produce sexuales which lay over-
wintering eggs on the bark.
Overwintering populations on crucifer roots identified as P. populitrans-
versus give rise to another wave of sexuparae in spring. These sexuparae also
produce sexuales on cottonwood, the eggs of which hatch later than the over-
wintering eggs, so that there are two discrete times of gall initiation. Galls on
early, proximal leaves are more elongate, those on later, distal leaves are more
globular (Bird et al., 1979; Rhomberg, 1980). From recent studies in Arizona
(Moran, 1993; Aoki and Moran, 1994), it seems probable that populations
producing the globular galls are a distinct species, P. obesinymphae, which
has specialized first instar gall defenders like P. spyrothecae. Both types of
gall occur throughout most of the range of Populus deltoides (Bird et al.,
1979). Geographic variation of this species complex has been studied exten-
sively using multivariate methods (Sokal and Riska, 1981). Bingham and Sokal
(1986) studied host tree effects on morphology.
Pemphigus populitransversus is also nominally recorded on Populus sp.
in Mexico (BMNH colln, leg. R. Peña), Argentina (Chiesa Molinari, 1942) and
Chile (BMNH colln, leg. G. Monsalve); also South Africa (Müller and Scholl,
1958), where it occurs on Populus nigra (BMNH colln, leg. E. McC. Callan)
and P. deltoides, and the Azores (on Populus sp.; BMNH colln, leg. A.
van Harten). Presumed anholocyclic populations occur in New Zealand on
Eutrema wasabi roots (BMNH colln, leg. L.M. Emms); aphids collected on
roots of Raphanus raphanistrum in Parana, Brazil (VFE) and on Arabis roots
in England (BMNH colln, leg. G. Fox-Wilson) may also be this species.
2n = 20.

Pemphigus populivenae Fitch Galls are elongate pocket-shaped or cocks-

comb-shaped, greenish or yellowish or pale red, arising alongside mid-rib on
upper side of leaf, with a slit-shaped opening underneath. On heavily-infested
trees, galls may occur on both sides of the leaf and those on the under-
side may then be confused with those of P. betae (Maxson and Knowlton,
1929). On cottonwoods (angustifolia, balsamifera, deltoides, fremontii and
trichocarpa; the latter in experimental conditions only). Fundatrix yellowish-
 Pemphigus 811

green. Alatae (BL 1.1-1.8 mm) emerge in June-July and migrate to found
colonies on roots of Chenopodiaceae. The evidence for this host alterna-
tion is work in California by Grigarick and Lange (1962), who transferred
sexuparae from sugarbeet to poplar bark under controlled conditions and
obtained fundatrices that produced galls specifically on P. trichocarpa. The
alatae from these galls were identified as P. populivenae, and colonized roots
of B. vulgaris and Chenopodium album. However, the suggested synonymy
of both P. betae and P. balsamiferae with populivenae indicates some confu-
sion of identity and further work seems advisable to confirm that the correct
name for populations damaging sugarbeet in California (e.g. Summers and
Newton, 1989) is in fact populivenae rather than betae. There may be a
complex of species with different Populus spp. as primary hosts. Whitham
(1978) reported that P. populivenae galling Populus fremontii in Utah caused
chlorosis of the leaf distal to the gall.

Pemphigus protospirae Lichtenstein Galls on Populus spp., especially

P. nigra var. italica, are smooth, shiny, green mottled with red, caused by
swelling, flattening and spiral twisting of the leaf petiole, similar to those
of P. spyrothecae but usually thinner and with more spirals (Fig. 131R).
Pemphigus protospirae, however, is heteroecious, and the alatae leave the gall
in late May-early July to found colonies usually in the sheathing leaf-bases
of aquatic Umbelliferae (Berula erecta, Sium latifolium, Apium nodiflorum;
Stroyan, 1964b; Heie, 1980). Widely distributed in Europe, but probably
restricted by availability of secondary host habitats. Also recorded from
western Siberia and Central Asia, but records from further east than this
(e.g. China; Chang and Zhong, 1979c) are doubtful.

Pemphigus saliciradicis (Börner) Apterae yellowish-white, lightly coated

with greyish wax; BL 1.6-2.4 mm. On roots of Salix spp. (fragilis, herbacea,
polaris) in Europe (records are from England, Finland, Sweden, Switzerland),
Greenland (as salicicola Hille Ris Lambers) and Japan. Aphids on Salix roots
in California, USA, and on Baffin Island in arctic Canada, are possibly also
this species (see below). Palaearctic populations are apparently normally
anholocyclic on Salix roots; alatae are unknown from Europe, but inter-
mediates between virginoparae and sexuparae occur in Japan (Aoki, 1975).
Lange (1965) reported Pemphigus populations of the saliciradicis group on
Salix roots in northern and southern California; the southern population
occurred throughout the year on Salix roots, whereas the northern population
produced sexuparae in autumn. However, no descriptions were published of
the Californian forms. Richards' (1963) record of P. groenlandicus (a grass-
root feeder) from roots of Salix reticulata on Baffin Island may be referable
to the saliciradicis group.

Pemphigus sinobursarius Chang Galls on P. simonii in China are on under-

sides of leaves at base of the lamina (Chang and Zhong, 1979c). Alatae similar
to P. bursarius, but with a shorter R IV+V (0.065 mm). Heteroecious, secon-
dary host unknown.
812 Pemphigus

Pemphigus siphunculatus Hille Ris Lambers Galls on twigs of Populus

ciliata in Pakistan in August contained apterous progeny of the fundatrix,
colour in life unknown (presumably wax-covered); BL 1.9-3.4 mm (Hille Ris
Lambers, 1973b). Presence of apterae in the gall is normally indicative of a
monoecious life cycle but, according to S. Chakrabarti (pers. comm. cited by
Aoki and Kurosu, 1991d), P. siphunculatus is heteroecious. The secondary
host is not recorded.

Pemphigus spyrothecae Passerini Gall is green, reddish or yellowish,

smooth, formed by thickening, flattening and spiral twisting of petiole of
Populus nigra (Fig. 131S). Gall formation was studied by Dunn (1960).
Fundatrix is pale green, giving rise to second generation apterae within the
gall. Monoecious holocyclic; alate sexuparae emerge in August-November, to
produce sexuales on the bark. Lampel (1960) provided a detailed account.
First instars with thick fore legs function as soldiers and defend the gall
against predators (Aoki and Kurosu, 1986; Foster, 1990). Widely distributed
in Europe, in North Africa (Tunisia), western Siberia, and introduced into
western Canada. 2n = 20.

Pemphigus tibetensis Chang Galls are globular, pouch-like, formed on twigs

of Populus cathayana in Tibet (Chang and Zhong, 1979c). This appears to
be a member of the borealis/dorocola group.

Pemphigus trehernei Foster Galls induced experimentally on Populus nigra

var. italica were green with reddish tinge around exit hole, globular, formed
at base of current year's shoot. Fundatrix is dull pale green. Alatae (BL
1.9-2.4 mm) emerge in June-July and found colonies on roots of Aster
tripolium in salt-marsh habitats (Foster, 1975). In England, Ireland, France.
Anholocyclic overwintering on Aster roots is common and the species has only
been found naturally on the secondary host; the gall generations were obtained
by experimental transfer of sexuparae to P. nigra.

Pemphigus venosus Habib and Ghani Recorded as forming galls on twigs

of Populus ciliata in Pakistan, but no description is given, and therefore this
is a nomen nudum.

Pemphigus vesicarius Passerini Galls on Populus nigra originate from mid-

rib at base of upper side of leaf, developing in April-May into an irregular
pale green structure with numerous tubular outgrowths, of diameter up to
4cm, that encompasses the whole leaf (Fig. 131T). Fundatrix dark slate
grey to blue-black. Alatae, BL 2.0-2.5 mm, leave gall in May-June through
secondary holes at apices of outgrowths (Roberti, 1938). Heteroecious;
secondary host unknown. In southern Europe, southwest and Central Asia.
 Periphyllus 813

PERIPHYLLUS van der Hoeven Chaitophorinae

About 40 species of medium-sized to large, generally long-haired aphids, living

on Acer except for a few species on Aesculus and one on Koelreuteria. The
antennal terminal process is usually long and the siphunculi are stump-shaped,
usually with polygonal reticulation, but often longer than in Chaitophorus and
with a pronounced flange. CAUDA is either rounded or tongue-shaped with
a slight constriction. Colonies are usually ant-attended. The association with
the host plant is an intimate one, with a complex seasonal polymorphism to
cope with physiological changes in the host, the most unusual feature of which
is the frequent occurrence of specialized larval aestivating morphs (Fig. 5,
p. 11). Periphyllus is predominantly a palaearctic genus (15 European, 9
northern Indian/Himalayan, 13 East Asian), with just three species native to
North America. Essig and Abernathy (1952) wrote a monograph on the genus.
There are keys to Periphyllus species of Europe (Hille Ris Lambers, 1966a),
Britain (Stroyan, 1977), Fennoscandia and Denmark (Heie, 1982), Ukraine
(Mamontova, 1955), India (A.K. Ghosh, 1980; Chakrabarti et al., 1987),
China (Tao, 1964), eastern Siberia (Pashchenko, 1988a, b), Japan (Sorin,
1990b) and Canada (Richards, 1972a). Aphidius aereolatus, A. setiger, Praon
negundinis and Trioxys falcatus are specialized parasitoids.

Periphyllus acericola (Walker) Apterae pale green or yellowish-green, some-

times with dorsal brownish markings; BL 2.4-3.5 mm. Alatae have broad dark
dorsal abdominal cross-bars scarcely separated between segments and paler
marginal sclerites; BL 3.0-3.5 mm. On undersides of leaves, petioles and
young shoots of Acer pseudoplatanus throughout most of Europe. Aestivating
first instar nyrnphs are yellowish-white with long, pointed dorsal and marginal
hairs; they aggregate in dense groups appearing like whitish spots on under-
sides of leaves. Monoecious holocyclic; oviparae and alate males in October.
2n = 18*.

Periphyllus acerihabitans Zhang Appearance in life unknown; BL of aptera

c. 2.5mm. ANT PT is unusually short; less than 2 x BASE VI. On Acer
buergerianum in China (Zhang and Zhong, 1982c). Biology unknown.

Periphyllus aceriphaga Chakrabarti and Mandal Apprearance in life

unknown; BL 2.4-2.7 mm. On Acer spp. (caesium, caudatum) in Uttar
Pradesh, India (Chakrabarti et al., 1987). Aestivating nymphs have foliate
marginal hairs. Life cycle and sexual morphs unknown.

Periphyllus aceris (Linnaeus) Apterae yellow, often with green flecks dor-
sally; BL 1.5-3.7 mm. Alatae have dorsal cross-bands more widely separated
than in P. acericola, with equally dark marginal sclerites and pterostigma; BL
814 Periphyllus

3.2-4.5 mm. On undersides of leaves, petioles and growing shoots of Acer

spp., especially A. platanoides, which is clearly its normal host. There are also
records from Aesculus, but no authentic specimens have been seen and these
should possibly be referred to P. testudinaceus. Throughout most of Europe,
and introduced to North America. Aestivating nymphs have long pointed
dorsal and marginal hairs and aggregate like those of P. acericola. Monoe-
cious holocyclic; oviparae and alate males in October-November. This species
has been confused in the literature with acericola and also with other species;
e.g. Börner (1940) had P. lyropictus and Wahlgren (1935) had P. testudina-
ceus. 2n = 16*.

Periphyllus aesculi Hille Ris Lambers Apterae rather elongate oval, brownish
in life with black dorsal spots (A.K. Ghosh, 1980); BL 2.5-3.5 mm. Siphunculi
wholly dark. Alatae have short dorsal abdominal cross-bars and dark marginal
sclerites. On undersides of young leaves of Aesculus indicus in India (Kashmir,
Himachal Pradesh, Uttar Pradesh) and Pakistan (BMNH colln, leg. CIBC).
Chakrabarti et al. (1972a) described the apterous vivipara and redescribed the
alate vivipara; L.K. Ghosh (1986) described the ovipara; and Agarwala and
D.N. Raychaudhuri (1982) described the alate male. Aestivating morph not
recorded. Sexuales in October-December.

Periphyllus allogenes Szelegiewicz Apterae elongate oval, brown with dark

brown antennae and legs; BL 1.0-1.2 mm. Alatae unknown. On leaf petioles
of Acer triflorum in Korea. Described from autumnal (late August) apterae,
presumed to have developed from aestivating nymphs and very different from
spring forms, which are probably much larger (Szelegiewicz, 1981). Biology
unknown.

Periphyllus americanus Baker Apterae in life 'pale yellowish-green with

dark median stripe on abdomen forming a cross in region of cornicles'
(Hottes, quoted by Palmer, 1952); BL 2.0-2.5 mm. Alata dark greenish
with black dorsal abdominal cross-bars and marginal sclerites. On leaves of
Acer spp., especially sugar maples, widely distributed in North America.
Aestivating nymphs have foliate marginal hairs. Monoecious holocyclic;
Essig and Abernathy (1952) gave an account of the summer generations,
but fundatrices and sexuales have apparently not been described. However,
there is an alate male in the BMNH colln (leg. G.F. Knowlton), from Utah
in October.

Periphyllus bengalensis Ghosh and Raychaudhuri Apterae yellow, with

dark siphunculi, pear-shaped; BL 1.8-2.2 mm. Alatae with dark dorsal
abdominal cross-bands. Forming a colony on undersides of older leaves of an
unidentified Acer sp. in West Bengal, India (A.K. Ghosh, 1980). Aestivating
and sexual morphs unknown.

Periphyllus brevisetosus Sorin Appearance in life unknown; only late sum-

mer apterae (BL 1.2-1.3 mm) are described, collected in Japan on Acer
 Periphyllus 815

'picutum' (pictum?) var. typicum (= mono) (Sorin, 1990b). Closely related to

P. kuwanii.

Periphyllus brevispinosus Gillette and Palmer Apterae brown with dark and
pale mottling, and pale siphunculi; BL 2.4-2.9 mm. Alatae brown with black
dorsal abdominal cross-bands and marginal sclerites; on undersides of leaves
of Acer glabrum in western North America. Aestivating nymphs have foliate
marginal hairs with serrate margins (Essig and Abernathy, 1952). Monoecious
holocyclic; oviparae in late September (Palmer, 1952).

Periphyllus bulgaricus Tashev Apterae shining black, with pale legs and
antennae; BL 2.0-2.8 mm. Apterae and alatae both have broad black dorsal
abdominal cross-bars. On shoot tips and undersides of leaves, usually at
base of mid-rib. Described from Acer hyrcanum in Bulgaria (Tashev, 1964);
also collected from A. hyrcanum in Turkey, from A. opulifolium in Iran,
from A. hermaneum in Lebanon (BMNH colln, leg. DHRL) and from
A. granatense and A. monspessulanum in Spain. Monoecious holocyclic;
Nieto Nafria and Gonzales Funes (1990) described fundatrices, oviparae and
alate males and discussed the very difficult separation of this species from
P. rhenanus. Aestivating nymphs not recorded.

Periphyllus caesium Chakrabarti and Saha Appearance in life not recorded,

probably rather pale with dark siphunculi; BL of aptera 2.3-2.4 mm. Alatae
have dark dorsal abdominal cross-bands. On Acer caesium in Uttar Pradesh,
India (Chakrabarti et al., 1987). An alate male ascribed to this species was
collected in October. Longer haired than P. aceriphaga and with a shorter HT
II, but otherwise apparently very similar.

Periphyllus californiensis (Shinji) Apterae dark olive green to brown, with

dark brown head and prothorax and irregular dark dorsal segmental spots
or cross-bands; BL 2.3-3.5 mm. Alatae have dorsal abdominal cross-bands
which, with a hand lens, appear darker than the pterostigma of the fore-
wing (cf. aceris). On Acer spp. in East Asia (China, Japan, Siberia, Korea,
Taiwan, Thailand, West Bengal), living on undersides of leaves along veins;
Pashchenko (1987) gave A. pseudosieboldianum as the principal food plant.
Also recorded from Aesculus turbinata in Japan (as acerifoliae; Takahashi,
1919a). Introduced to Europe, North America, Australia and New Zealand,
where it is most commonly found on planted Asian ornamental maples, but
sometimes also colonizes native species; there is one record from Aesculus
californica in California. Aestivating nymphs have foliate marginal hairs.
Monoecious holocyclic; Essig and Abernathy (1952) fully described the life
cycle in California, where oviparae and alate males occur in October-
November. Population dynamics of P. californiensis in Japan has been
studied in relation to host phenology (Hashimoto and Furuta, 1988), mortality
of aestivating nymphs (Furuta, 1985), host preferences (Furuta, 1986) and
natural enemies (Furuta and Mao, 1986; Kan and Sasakawa, 1986). 2n = 18
(Blackman, 1986).
816 Periphyllus

Periphyllus coracinus (Koch) Apterae variable in colour, yellow, light or

dark green, or brown, or mottled with green or brown. On leaf petioles
and twigs of Acer platanoides in Western, Central and Eastern Europe (The
Netherlands, Sweden, Germany, Austria, Poland, Hungary, Ukraine).
Monoecious holocyclic; oviparae and alate males in October. Apparently no
specialized aestivating nymphs are produced.

Periphyllus formosanus Takahashi Apterae are reddish-brown, with thorax

darker; BL c. 2.5 mm. Alatae have blackish dorsal abdominal cross-bands.
On undersides of leaves of unidentified Acer sp(p). in Taiwan and China.
Aestivating nymphs have foliate marginal hairs (Takahashi, 1933a). Monoe-
cious holocyclic; probable fundatrices were observed by Takahashi in
February, and Essig and Abernathy (1952) described the oviparae and alate
males, collected in November.

Periphyllus garwhalensis Chakrabarti and Mandal Apterae are undescribed.

Alatae (BL 3.0-3.6 mm) have dark siphunculi but no dorsal abdominal cross-
bands. On Acer caudatum in Uttar Pradesh, India. Aestivating nymphs have
foliate marginal hairs with serrate margins (Chakrabarti et al., 1987).

Periphyllus himalayensis Chakrabarti Apterae undescribed. Alatae with

dark dorsal abdominal cross-bands, colour in life unknown, BL 3.1-3.5 mm
(Chakrabarti, 1977). On unidentified Acer sp(p). in northern India (Himachal
Pradesh, West Bengal). Biology unknown. Perhaps a synonym of P.
bengalensis.

Periphyllus hirticornis (Walker) Apterae pale green, without dark markings;

BL 2.0-3.0 mm. Abdomen of alata with variably developed spinal sclerites,
but not cross-bands. On undersides of leaves, leaf petioles and fruits of
Acer campestre in Europe, often with attendant ants. Aestivating nymphs
have foliate marginal hairs (see Stroyan, 1977). Monoecious holocyclic;
oviparae and alate males in October-November. 2n = 18*.

Periphyllus hokkaidensis Sorin Apterae dark reddish to blackish-brown;

BL c. 2mm. On Acer miyabei and A. mono var. subtrifidum in Hokkaido,
Japan (Sorin, 1990b). Other morphs and biology unknown. Closely related to
P. kuwanaii.

Periphyllus koelreuteriae (Takahashi) Apterae variably pigmented, yellow

or black, with wholly black tibiae, and often with paired dark dorsal spots
on abdominal tergites; BL 1.6-2.5 mm. Alatae have dark dorsal abdominal
cross-bars, and 25-38 secondary rhinaria on ANT III (cf. alatae of califor-
niensis, with 7-25 secondary rhinaria on III). On Koelreuteria paniculata in
China, Korea and Japan. Aestivating nymphs have foliate marginal hairs
(Takahashi, 1933a; Essig and Abernathy, 1952). The recorded occurrence
on Aesculus turbinata is apparently casual, being based on a single alata
(Higuchi, 1972). Sorin's (1990b) apterae from A. turbinata keyed as P.
 Periphyllus 817

koelreuteriae do not appear to be this species. Sexuales not described.

2n = 18*.

Periphyllus kuwanaii (Takahashi) Apterae and alatae shining black; BL

1.4-2.4 mm. Principally on Acer mono; Pashchenko (1988a, b) recorded it
from several other Acer spp., but such visits may be casual. In China, Siberia,
Japan, Korea. Aestivating nymphs have foliate marginal hairs (Takahashi,
1933a); but part of the population does not aestivate (Pashchenko, 1988b).
Sexual morphs apparently undescribed. Periphyllus diacerivorus Zhang (in
Zhang and Zhong, 1982c) seems to be a synonym. 2n = 18*.

Periphyllus loricatus Pashchenko (= ginnalae Paik in Szelegiewicz, 1974)

Apterae and alatae shining dark brown to black; BL 1.6-2.5 mm. On Acer
ginnale, living in early spring on leaf petioles at tips of young branches, and
later forming dense colonies on undersides of leaves. Aestivating nymphs are
green to brown with foliate marginal hairs, living singly on both sides of
leaves; but most of the population apparently does not aestivate (Pashchenko,
1988b). In Siberia, Japan and Korea. Monoecious holocyclic; apterous males
and oviparae in September (Pashchenko, 1987). Periphyllus takahashii Sorin,
1990b, described from A. ginnale in Japan, appears to be a synonym.

Periphyllus lyropictus (Kessler) Apterae yellowish with brown dorsal mark-

ings, usually comprising a broad spinal stripe on head and thorax and a large
V-shaped mark on dorsal abdomen; BL 1.9-3.0 mm. Alata have dark marginal
sclerites, but other dorsal sclerotization is limited to the spinal area, not
forming cross-bands. On undersides of leaves of Acer platanoides, often
forming large colonies producing much honeydew and visited by ants and
other insects. This species does not produce aestivating nymphs. In Europe,
and introduced on Norway maple to North America where it is also wide-
spread. Acer platanoides is clearly the favoured host and records from other
Acer are mostly due to confusion with P. aceris and P. testudinaceus,
particularly in the North American literature (see Essig and Abernathy, 1952).
Monoecious holocyclic; oviparae and alate males in October-November.

Periphyllus mamontovae Narzikulov Apterae green; BL 2.1-2.9 mm. On

leaves of Acer turkestanicum and (more rarely) A. regelii in Tadzhikistan
(Narzikulov, 1957). Aestivating nymphs are apparently not produced. Life
cycle and sexuales are undescribed.

Periphyllus mandshuricus Pashchenko Apterae apparently not produced

in spring; reddish-brown to dark brown fundatrices on shoot tips produce
only dark brown alatae (BL 2.5-3.4 mm), which live on undersides of leaves
and produce aestivating nymphs with foliate marginal hairs. On Acer mand-
shuricum in maritime region of Siberia. Oviparae and alate males in October
(Pashchenko, 1988b).
Periphyllus minutus Shaposhnikov Apterae shining brown or greenish-
yellow, with dark brown dorsal markings; BL 1.6-2.5 mm. Alatae have dark
818 Periphyllus

dorsal abdominal cross-bands. On Acer tataricum, colonizing young shoots,

leaf petioles, undersides of leaves, fruit stalks and fruits. Aestivating nymphs
have long pointed dorsal and marginal hairs. In Eastern Europe and south-
west Asia (Bulgaria, Hungary, Iran, Kazakhstan, Ukraine). Sexuales not
described. Described as a subspecies of aceris by Shaposhnikov (1952);
Mamontova (1955) provided a fuller description and gave it full species
status.

Periphyllus montanus Sorin Colour in life unrecorded; BL of aptera

c. 1.6mm. Alata has broad and dark dorsal abdominal cross-bands. On Acer
rufinerve in Japan (Sorin, 1979b). Biology and sexual morphs are unknown.

Periphyllus negundinis (Thomas) Apterae pale yellow-green to apple green

with dark dorsal markings; BL 2.0-2.5 mm. Alatae have variable dark green
dorsal abdominal markings, but no distinct cross-bands. On Acer negundo,
feeding on young growth in or near fruit clusters in spring, then on undersides
of leaves. Aestivating nymphs have foliate marginal hairs. Monoecious
holocyclic; apterous males and oviparae in October-November. Essig and
Abernathy (1952) provided descriptions of morphs and life cycle. 2n = 20.

Periphyllus nevskyi Mamontova Apterae shining grey-black, with a central

black sclerite extending across ABD TERG 4-6; BL 1.3-2.0 mm (Nevsky
1929a, as Chaitophorinella negundinis; Mamontova, 1955). On Acer regelii in
Central Asia (Tadzhikistan, Uzbekistan). Also recorded from A. turkestanica
and A. pseudoplatanus (Gabrid, 1989). Life cycle not studied.

Periphyllus obscurus Mamontova Apterae blackish-green; BL 1.8-2.6 mm.

In ant-attended colonies on young shoots, leaf petioles and undersides of leaves
of Acer campestre in Europe (Bulgaria, the former Czechoslovakia, England,
The Netherlands, Germany, Hungary, Poland, Turkey, Ukraine). Aestivating
nymphs are apparently not produced. Sexual morphs not described.

Periphyllus pallidus Chakrabarti and Saha Appearance in life not recorded;

BL of aptera 2.4-2.7 mm. On an unidentified Acer sp. in Uttar Pradesh, India
(Chakrabarti et al., 1987). Aestivating nymphs have foliate marginal hairs.
Other morphs and life cycle unknown.

Periphyllus rhenanus (Börner) Apterae shining, dark; BL 2.5-3.0 mm. On

undersides of leaves, extending onto petioles and twigs. Alatae have broad
black dorsal abdominal cross-bars and equally dark marginal sclerites. On
Acer monspessulanum and A. obtusatum in southern and Central Europe
(Bulgaria, France, Germany, Greece, Italy, Switzerland). Aestivating nymphs
have very long marginal hairs and form dense clusters (Barbagallo and
Stroyan, 1982). Monoecious holocyclic; apterous males and oviparae occur
in October-November (Hille Ris Lambers, 1947c; as P. lichtensteini). Nieto
Nafria and Gonzales Funes (1990) discussed differences from the closely-
related P. bulgaricus.
 Periphyllus 819

Periphyllus singeri (Börner) Apterae reddish-brown, in spring bearing dark

dorsal abdominal cross-bands like the alatae; BL 2.4-3.5 mm. On young
shoots of Acer pseudoplatanus, attended by ants. In Western and Central
Europe (Austria, Germany, The Netherlands). Aestivating nymphs are
apparently not produced. Monoecious holocyclic; oviparae and alate males in
October (BMNH colln, leg. DHRL).

Periphyllus steveni Mamontova Apterae elongate oval, shining brown or

dark green; BL 1.8-2.4 mm. On young growth and leaves of Acer stevenii
in Crimea (Mamontova-Salukha, 1962). One of the group of species with long
black dorsal hairs (Nieto Nafria and Gonzales Funes, 1990). Aestivating
nymphs are yellow, with long pointed hairs. Life cycle and sexual morphs not
described.
Periphyllus tegmentosus Pashchenko Fundatrix green with brown markings;
her progeny are all alate, having shining greenish-yellow abdomen with dark
dorsal abdominal cross-bars and marginal sclerites. BL of alata 2.2-2.9 mm.
On Acer tegmentosum in maritime Siberia. Aestivating nymphs yellow-green
with foliate marginal hairs. Monoecious holocyclic; oviparae and alate males
in October (Pashchenko, 1988b).

Periphyllus testudinaceus (Ferni), Plate l1c, d Apterae dirty dark green to

dark brown or blackish; BL 2.0-3.7 mm. Alatae have dark dorsal abdom-
inal cross-bands, darker than pterostigma, and equally pigmented marginal
sclerites. On young growth, leaves and leaf petioles of numerous Acer spp.
(especially on A. campestre and A. pseudoplatanus) and sometimes on
Aesculus spp. Throughout Europe, in Korea (needs confirmation) and intro-
duced to Tasmania, New Zealand and North America - where it is widely
distributed. Aestivating nymphs with foliate marginal hairs are the only morph
occurring from June to September (Fig. 5). Oviparae and alate males in
October-November. Essig and Abernathy (1952) provided a detailed account
of the life cycle and polymorphism. Warrington and Whittaker (1985) showed
that ant attendance caused a large increase in sap ingestion by this species.
2n = 18.
Periphyllus tokyoensis Sorin Appearance in life not recorded. BL of
aptera only c. 1.1 mm. On an unidentified Acer sp. in Japan (Sorin, 1990b).
Aestivating nymphs have foliate marginal hairs. Other morphs and life cycle
unknown. Possibly these are dwarf specimens of a previously known species.
Periphyllus vandenboschi Hille Ris Lambers Colour of aptera in life not
recorded but probably pale green, with contrasting black siphunculi; BL
1.4-2.0 mm. Alata has pale brown abdomen with dark markings only on ABD
TERG 4-8 (Verma, 1974). Collected on leaf petioles of an unidentified Acer
in Pakistan (Hille Ris Lambers, 1966c) and also recorded from Himachal
Pradesh, India (see L.K. Ghosh, 1986). Aestivating nymphs and other
morphs, including sexuales, have not been recorded. [Chakrabarti et al.
(1972a) had P. villosii.]
820 Phloeomyzus

Periphyllus venetianus Hille Ris Lambers Apterae shining blackish-green

with pale antennae and legs; BL 1.1-2.0 mm. Alatae are undescribed. On
undersides of leaves and leaf petioles of Acer campestre, attended by ants.
Aestivating nymphs have foliate marginal hairs. In southern Europe (Austria,
former Czechoslovakia, France, Italy, former Yugoslavia). Monoecious holo-
cyclic; Barbagallo and Stroyan (1982) recorded immature oviparae in Sicily in
September.

Periphyllus villosii Chakrabarti Aptera elongate oval, large, probably pale;

BL c. 3.5mm. Abdomen of alata without any dark dorsal cross-bands. On
Acer spp. (caesium, caudatum, villosum) in Uttar Pradesh, India. Aestivating
nymphs have foliate marginal hairs (Chakrabarti et al., 1987). Oviparae and
alate males (on A. villosum) in October (Chakrabarti, 1977).

Periphyllus viridis (Matsumura) Apterae (?fundatrices) green; BL 2.9-

3.9mm. Alatae (in spring) without dorsal abdominal cross-bands; according
to Pashchenko (1987), all progeny of the fundatrix are alate. On young
growth, leaves and leaf petioles of Acer mono in Japan, Korea and maritime
Siberia. Aestivating nymphs have foliate marginal hairs. Oviparae and alate
males in October; Pashchenko (1987) gave a full account of the life cycle.
Sorin (1990b) described some large alate specimens in Japan as a subspecies,
P. viridis osugiensis.

PHLOEOMYZUS Horvath Phloeomyzinae

Perhaps only one species, unique in morphology and in that the only alatae
are sexuales. Parthenogenetic forms are all apterous, with fused head and
prothorax and 3-faceted eyes.
Phloeomyzus passerinii Signoret, Plate 2e, f Apterae green, covered with
dirty white wax wool; BL 1.2-2.2 mm. On bark and in crevices on trunks of
Populus spp. (alba, ciliata, x euroamericana, maximowiczii, nigra, suaveolens,
tadzhikistanica, tremuloides), heaviest infestations occurring on six to eight-
year-old trees. Monoecious holocyclic; oviparae and males are produced in
September-October in the northern hemisphere, the oviparae each laying two
eggs (Theobald, 1929). Anholocyclic overwintering by apterous viviparae is
also common and may predominate; no morph recognizable as a fundatrix
has been described. In Europe, North Africa (Egypt, Morocco), southwest,
Central and East Asia (incl. Japan; BMNH colln, leg. S. Aoki); USA (only
record is from Maine) and South America (Argentina, Chile, Colombia).
Arzone and Vidano (1984) studied the life cycle in Italy. The European
P. nigra-feeding form was described as a separate species, Ph. redelei Hille
Ris Lambers, 1931 (see Doom and Hille Ris Lambers, 1962), as also were
large specimens found on P. tremuloides in Maine (as Ph. dearborni; Smith,
 Phyllaphis 821

1974b). Zhang (in Zhang and Zhong, 1982b) described a subspecies in China,
Ph. passerinii zhangwuensis. It seems likely, however, that a single, somewhat
variable species is involved. 2n = 10.

PHORODON Passerini Aphidinae: Macrosiphini

About four Myzus-like species with finger-like processes on antennal tubercles

and a secondary host association with Cannabidaceae (Cannabis, Humulus).
The heteroecious species have Prunus as primary hosts. The record of
Ph. humulifoliae from Ulmus sp. (Tao, 1963) is probably in error. See also
B & E, 1984, p. 327.

Phorodon humuli (Schrank) Apterae are pale green to yellowish-green,

rather shiny, with three darker green dorsal longitudinal stripes; BL 2.0-
2.6mm. Alatae have a black dorsal abdominal patch. Spring colonies occur
on Prunus spp. (domestica, insititia, mahaleb, spinosa), causing only slight
leaf curl. It is a vector of plum pox virus. In Europe, North Africa (Ethiopia),
southwest Asia (Iran, Turkey); introduced to North America, where it is
widespread, and New Zealand (BMNH colln, leg. P.L. Robertson). Hetero-
ecious holocyclic; migration occurs in May-June to hops (Humulus lupulus),
on which its importance as a pest has increased with development of insecticide
resistance (Hrdy et al., 1986). Return migration to Prunus (studied by Eppler,
1988) is in September-October. Morphometric variation was studied by
Hampson and Madge (1986a, b) and allozyme variation by Eggers-Schumacher
and Sander (1988). See also B & E, 1984, p. 328. 2n = 12.

Phorodon japonensis Takahashi Apterae and alatae resemble those of

Ph. humuli (Takahashi, 1965b). On Prunus spp. (mume, salicina) in Japan,
Korea, eastern Siberia and Taiwan. Heteroecious holocyclic; migration to
Humulus spp. (japonicus, scandens). Described as a subspecies of humuli, to
which it is closely related. 2n = 12.

PHYLLAPHIS Koch Drepanosiphinae: Phyllaphidini

A genus of two or three species on Fagus with well-developed dorsal wax

glands, a very short antennal PT and pore-like siphunculi. Takahashi (1960b)
distinguished it from related genera with a key, Stroyan (1977) provided a
generic diagnosis and Richards (1973) keyed the apterous viviparae. Trioxys
phyllaphidis is a specialized parasitoid.
822 Phyllaphoides

Phyllaphis fagi (Linnaeus), Plate 8e, f Apterae elongate oval, pale yellowish-
green, covered with wax wool; BL usually 2.0-3.2 mm (but summer dwarfs
down to 1.1 mm). Alatae have abdomen wax-covered, with variably-developed
dark dorsal cross-bars. On undersides of mostly young leaves of Fagus
spp., causing them to curl downwards on both sides of the mid-rib, and often
to wither and die prematurely. Throughout Europe, east to Turkey and
Caucasus, and introduced to Australia, New Zealand and North America.
Records from Japan apply to Ph. fagifoliae. Monoecious holocyclic, with
dwarf apterae living through summer months to produce apterous oviparae
and alate males in October-November. Oviparae have dark dorsal cross-
bars and dark siphunculi (cf. Ph. grandifoliae). Chandler (1968) studied its
association with a syrphid predator. 2n = 16.

Phyllaphis fagifoliae Takahashi Apterae pale, presumably with wax; BL

1.2-2.5 mm. Differing from Ph. fagi in having much less pigmented alatae
(Takahashi, 1937c). On Fagus spp. (crenata, japonica) in Japan. Monoecious
holocyclic; sexuales in October (Higuchi, 1972; as Ph. fagi). 2n = 26 (after
Shinji; see Blackman, 1986).

Phyllaphis grandifoliae Richards Apterae pale greenish, pale yellow or

somewhat cream-coloured (Richards, 1973), presumably with wax; BL
c. 1.8mm. On Fagus grandifolia in western North America. Monoecious
holocyclic; oviparae have pale siphunculi and lack dorsal abdominal cross-
bars (Smith and Parron, 1978). The distinction from Ph. fagi requires
confirmation.

Phyllaphis nigra Ashmead Apterae shiny black; BL c. 1.3mm. On tender

shoots of Quercus phellos var. laurifoliae (= Q. laurifolia) in Florida, USA
(Ashmead, 1881). Not recorded since original description. Probably not
belonging in this genus; perhaps a Lachnochaitophorus. Life cycle not known.

PHYLLAPHOIDES Takahashi Drepanosiphinae: Phyllaphidini

A genus for one East Asian narrow-bodied species living on bamboos.

Phyllaphoides bambusicola Takahashi Alatae white with pale yellow meso-

thorax, secreting white cottony wax; BL 1.5-2.5 mm. Apterae are also
reported to occur (Takahashi, 1923), but do not seem to have been described.
On undersides of leaves of Bambusa stenostachya (Takahashi, 1923) and
Phyllostachys spp. (Liao, 1976) in Taiwan and China (Tao, 1964). No sexual
morphs described; in Taiwan it is anholocyclic (Takahashi, 1923). There are
conflicting reports as to whether this species does (Liao, 1976), or does not
(Takahashi, 1921), jump when disturbed.
 Phylloxera 823

PHYLLOXERA Boyer de Fonscolombe Phylloxeridae

A genus containing nominally about 60 small pear-shaped phylloxerids living

on Juglandaceae and/or Fagaceae. More than half the nominal species
were described from galls on Carya in North America (Shimer, 1869, as
Dactylosphaera; Pergande, 1904; Stoetzel and Tedders, 1981; Stoetzel, 1993).
Only the pecan-feeding species have been worked on taxonomically since
Pergande's studies and the status of many of the Carya gall-makers is therefore
uncertain. Although many species are clearly monoecious on either Carya or
Fagaceae, host alternation has now been demonstrated for at least two species
(Stoetzel, 1985b). Accounts of the oak- and chestnut-feeding species are
available for Europe (Heinze, 1962; Barson and Carter, 1972) and North
America (Duncan, 1922). It is likely that most if not all of the North American
Carya-feeders would be more correctly placed in Moritziella (Holman, 1974).
Grassi (1912) fully reviewed all information on the group then available and
Barbagallo and Binazzi (1991) reviewed the Italian oak-feeding species. Hedin
et al. (1985) studied the biochemistry of gall formation on pecan. Stoetzel
(1985a) described the development of the sexuales, through a rapid series of
moults of pupiform larvae.

Phylloxera bipunctata Lichtenstein Apterae yellow with two orange spots,

living solitarily on leaves of Quercus sp(p). in southern France and Switzerland
(Lichtenstein, 1874a). Alatae unknown, species not recognized since original
(brief) description.

Phylloxera caryaeavellana Riley Galls projecting underneath leaf, globular,

pale greenish, like a small hazelnut (5-10 mm diam.); above leaf flat or slightly
concave. Galls mature in May-June, emergent alatae have orange abdomen,
BL 0.9-1.2 mm (Pergande, 1904). On Carya tomentosa in Washington, DC.
Life cycle unknown.

Phylloxera caryaecaulis (Fitch) Galls on young twigs, on petioles or at bases

of leaflet main veins, sometimes in clusters; globular, diam. 5-25 mm, pale
yellowish-green tinted with red before opening, afterwards becoming leathery
and black. Typically on Carya glabra in eastern USA. Monoecious holocyclic;
alate sexuparae emerging from galls have dusky wings and in life there are
apparently distinctive dorsal prothoracic markings and a pair of orange or
blackish spots on the posterior abdomen; BL 0.8-1.8 mm (small alatae pro-
duce mostly males, larger ones mostly females). Morgan (1909b, 1912, 1915)
studied the cytogenetics and chromosome cycle of this species. Shimer (1869)
described three other closely-related phylloxerid species producing somewhat
different twig or petiole galls on other species of Carya, which may or may
not be synonymous with Ph. caryaecaulis: spinosa, producing galls covered
824 Phylloxera

with fleshy spines on Carya sp. in Illinois (the immature inhabitants of these
galls were described by Shimer as having long spiny tubercles); caryaemagna,
producing velvety galls on petioles or at base of mid-rib on C. amara
(= cordiformis) in Illinois and Missouri; and subelliptica, producing ellipsoid
mottled green galls on side of twig of C. alba ( = ovata) in Illinois (see also
Pergande, 1904). Caldwell and Schuder (1979) studied the life cycle of the
form on C. ovata in Indiana. Shimer (1869) reported on natural enemies of
galls of caryaemagna in Illinois. 2n = 8.
Phylloxera caryaefallax Riley Galls on leaves of Carya alba (= ovata) are
thin-walled, 1-5 mm diameter, on upper side of leaf and convex with a central
nipple-like opening below leaf. Alatae with dark-veined wings and orange-
yellow abdomen, BL 2.3-2.8 mm, emerge in May-early June (Pergande,
1904). In Missouri, Illinois and states bordering the Mississippi at least as far
south as Tennessee. Life cycle unknown. [Morgan's (1912, 1915) cytogenetic
study of 'Phylloxera fallax' cannot apply to this species, as Morgan's species
was monoecious, with fundatrices that produced apterous sexuparae; its
identity is still in doubt.] 2n = 12 (Steffan, 1968).
Phylloxera caryaefoliae Fitch Galls on leaves of Carya glabra project
above leaf as pale green slender cones with pale rings at their bases, c. 5 mm
diam., almost flat on underside of leaf with a central nipple. Alatae, with
distinctively marked prothorax and orange abdomen, BL 0.8-1.0 mm, emerge
from the upper side of the gall (Pergande, 1904) in early June. In northeastern
USA (New York, Illinois, Iowa, Virginia). Life cycle unknown. 2n = 8
(Steffan, 1968).
Phylloxera caryaeglobuli Walsh Galls according to original description are
spherical, on leaf lamina between veins, opening in June to release alatae
through a simple longitudinal slit on underside of leaf. On shagbark hickory
(= Carya ovata) in Illinois and Missouri. Pergande (1904) recorded the
available information, and suggested that hemispherica Shimer might be a
synonym. Life cycle unknown. 2n = 22 (Steffan, 1968).
Phylloxera caryaegummosa Riley Galls are spindle-shaped, suspended from
the underside of leaf of Carya alba (= ovata) by a slender petiole; they are
white or yellowish, pubescent and sticky. In Illinois and Missouri, USA.
Alatae undescribed and life cycle unknown.
Phylloxera caryaeren Riley Galls develop in clusters along leaf petioles and
mid-ribs of leaflets of Carya glabra; they are reniform, pale green, 2-15 mm
max. diam., placed transversely to axis of petiole, deeply divided by a trans-
verse slit and opening into two halves (rather like a bivalve mollusc) when
mature (Pergande, 1904). The phylloxerid itself is still undescribed, and the
life cycle is unknown. In Illinois and Missouri, USA.
Phylloxera caryaescissa Riley Galls are globular, diam. 8-14 mm, pale
greenish or dirty yellowish-green, clothed with short fine yellowish hairs,
 Phylloxera 825

projecting about equally on both sides of leaf lamina, opening underneath

when mature (mid-May to mid-June) by a transverse slit (Pergande, 1904).
Alatae have orange abdomen; BL 1.0-1.2 mm. On Carya tomentosa in eastern
USA (Florida, Washington, DC). Life cycle unknown.

Phylloxera caryaesemen Walsh Galls rounded, small (diam. 0.3-2.0 mm)

and very numerous, yellowish-green to brownish-yellow, rather tough and
leathery, projecting more above leaf than below, with a more-or-less depressed
centre above and opening (in July-early August) by a central nipple-like
opening underneath. No alatae are produced; according to Walsh (1868, cited
by Pergande, 1904), first instar larvae emerge to found new galls. On Carya
glabra in Illinois and Missouri, USA; Shimer (1869) also recorded it from
C. amara (= cordiformis) in Illinois.

Phylloxera caryaesepta (Shimer) Galls on Carya alba (= ovata) have conical

projections both above and below leaf, with nipple-like apices fringed with
filaments when mature, the structure on both sides of the leaf somewhat
resembling a miniature sea-anemone (diam. 5-12 mm). Colour of gall varies
from pale yellowish-green to red. Alatae with dark orange abdomen, BL
0.8-1.2 mm, emerge in late May-June from upper side of gall (Pergande,
1904). In Illinois and Missouri, USA. On Carya glabra further east (New
York, Virginia), the same or a closely-related species forms galls with a
narrower conical structure above and below leaf (Ph. caryaesepta "var.
perforans"; Pergande, 1904). Life cycle unknown.

Phylloxera caryaevenae Fitch Galls on Carya tomentosa and several other

Carya spp. are distinctive, usually pale yellowish-green, elongate folds along
the main veins on the underside of the leaflet, with corresponding ridges
or carinae above (Stoetzel, 1993). The opening is a slit on the underside.
Monoecious holocyclic. Galls are mostly dry and empty by mid-August, but
no alatae are known; the sexuales are produced from late July by apterous
sexuparae in free-living colonies in deep bark fissures on the trunk and
branches (Pergande, 1904). Widely distributed in eastern USA, from New
York to Florida and as far west as Illinois and Missouri.

Phylloxera castaneae (Haldeman) Apterae pale yellow, BL c. 1 mm, on both

sides of leaves, especially along the mid-ribs, of Castanea spp. (dentata,
mollissima, pumila, sativa) in northeastern USA (New York, Maryland,
Washington, DC, West Virginia). Infested leaves become much distorted and
desiccated. Heteroecious holocyclic; Stoetzel (1985b) demonstrated host alter-
nation in West Virginia from galls on Carya tomentosa. The galls and gall
generations have not yet been described; the galls were similar to those of
P. intermedia (M.B. Stoetzel, pers. comm.). Alatae from these galls gave rise
to large populations on Castanea spp., causing yellow discoloration along
the leaf veins. Pergande (1904) drew attention to the fact that most of the
immatures on the upper sides of the chestnut leaves were bright orange (as
opposed to pale yellow) and had longer tubercles than those on the undersides;
826 Phylloxera

his name spinifera is available for this form should it prove to be specifically
distinct.

Phylloxera coccinea (von Heyden) Fundatrices in spring cause crinkling and

discoloration of margins of young oak leaves. Later generations of apterae
(yellowish-orange, BL 0.8-1.2 mm) lay eggs in compact circles and cause
necrotic spots on undersides of leaves (see Börner and Heinze, 1957). Mono-
ecious holocyclic on Quercus spp. (especially robur) throughout most of con-
tinental Europe, and east to Kazakhstan. Both apterous and alate sexuparae
are produced (Börner and Heinze, 1957). Phylloxera rutila Dreyfus is a
synonym (Börner, 1952-3). New Zealand records apply to Moritziella corti-
calis (Sunde, 1974).

Phylloxera confusa (Grassi) Fundatrices feed from veins on undersides of

oak leaves, causing the lobe of the leaf distal to the feeding point to fold
inwards. Later generations of apterae (BL 1.5-2.0 mm) are yellowish to
orange-yellow and develop in small groups on young twigs, on leaf scars
or under loose bark (Grassi, 1912). Monoecious holocyclic; sexuparae are
apterous. On Quercus robur and Q. petraea, only recorded from central Italy.

Phylloxera conica (Shimer) Galls are almost globular, projecting on both

sides of the leaf, diam. 3-7 mm in plane of leaf, deep purplish-brown, with
a roughened or granular surface texture, becoming pubescent and covered
with a sticky secretion when mature. Alatae, BL 0.8-1.2 mm with a pale yellow
abdomen, emerge when the gall splits open underneath in June and pre-
sumably fly to an unknown secondary host. Originally described from Carya
alba (= ovata) in Illinois, USA (Shimer, 1869), but the description above is
of galls on C. amara (= cordiformis) in Missouri identified as this species
by Pergande (1904). Riley (1875) suggested synonymy with Ph. depressa, but
this does not accord with Pergande's interpretation of the species.

Phylloxera davidsoni Duncan Apterae light orange to yellowish-brown; BL

c. 0.7 mm. Immatures are pale yellow, alatae orange with black thorax. On
undersides of tomentous young leaves of Quercus engelmanni in California,
USA (Duncan, 1922). Life cycle unknown.

Phylloxera deplanata Pergande Galls on leaves of Carya tomentosa are cir-

cular, diam. 1-5 mm, slightly convex above leaf and more prominent beneath,
with an oval orifice fringed with fine hairs (Stoetzel, 1993). Colour is reddish
to yellowish or greenish-yellow above leaf and purplish to dull greenish-yellow
beneath. Monoecious holocyclic; fundatrices produce sexuales in the galls in
May, but some alatae may also be produced. Galls are empty, brown and dry
by late June (Pergande, 1904). Only known from the District of Columbia,
USA.

Phylloxera depressa (Shimer) Galls on leaves of Carya alba (= ovata)

in Illinois, USA are numerous, smooth, diam. 4-6 mm, convex and pale
 Phylloxera 827

yellowish-green above leaf, whiter beneath with a central nipple-like projec-

tion and a round opening. Alatae, BL c. 1 mm with a yellow abdomen, are
present in the gall in late June (Shimer, 1869). Life cycle unknown. According
to Riley (1875), conifera Shimer is a synonym. 2n = 6 (Steffan, 1968).

Phylloxera devastatrix Pergande Clusters of galls occur on young twigs, leaf

petioles or flower stalks of Carya illinoensis in southern and southeastern USA
(Arkansas, Louisiana, Mississippi, Oklahoma, Texas), often causing serious
economic damage. The galls are globular, of irregular shape, smooth, green
or yellowish-green with a reddish tinge before maturity, diam. 3-15 mm. They
open in May-early June by means of a transverse split into four or more broad
bracts, releasing alatae that immediately start to oviposit on neighbouring
twigs, etc. The eggs develop into dwarf sexuales that mate and lay over-
wintering eggs under bark or at the base of the tree. Monoecious holocyclic;
H. Baker (1935) described the life cycle in Louisiana and reported a high
incidence of parasitism of galls. Carpenter et al. (1979) and Calcote (1985)
studied varietal and clonal resistance to P. devastatrix in pecan, Neel and
Hedin (1985) studied its effects on nut production and Hedin et al. (1985)
studied biochemical aspects of gall formation. Mitchell et al. (1984) studied
the gall-associated fauna.

Phylloxera foaae Börner Monoecious holocyclic on deciduous oaks, with a

life cycle similar to that of Ph. coccinea, but Q. petraea is the usual host plant.
Fundatrices usually in small leaf-edge folds; subsequent generations of apterae
yellowish-orange, BL 0.7-0.9 mm, causing necrotic spots on undersides of
leaves. Sexuparae are alate. Closely related to Ph. coccinea; Heinze (1962)
provided some distinguishing characters. In Italy, Germany, Ukraine.

Phylloxera foveata (Shimer) Galls on leaves of Carya amara (= cordiformis)

are small (1.5-8.4 mm diam.), with a dimpled central depression above,
opening beneath leaf by a rounded downy orifice. Alatae were plentiful in
late June. In Illinois, USA (Shimer, 1869). The same leaves carried galls of
Ph. minima and Ph. caryaesemen.

Phylloxera foveola Pergande Galls on leaves of Carya glabra in Virginia,

USA; circular, yellow-green tinged with pink or red, 3-5 mm diam., convex
and rather prominent above leaf with a deep central dimple or depression
which is pinkish in colour, part below leaf convex or depressed with a small
conical central nipple. Alatae (BL c. 1.2mm, with slender, orange abdomen)
are mature in galls in late May and most had emerged before June, indicating
that it is probably heteroecious, but secondary host is unknown. Possibly a
synonym of Ph. foveata; the emergence time was the principal reason given
for separating the two (Pergande, 1904), but this might vary according to
weather conditions or host plant.

Phylloxera georgiana Pergande Galls occurred singly or in clusters on tender

twigs and petioles of an unidentified Carya sp., presumably at Georgiana,
828 Phylloxera

Florida (Pergande, 1904). They were similar to those of Ph. devastatrix; more-
or-less globular, with a short nipple, greenish-yellow (sometimes reddened on
one side), or wholly dark red, 5-10 mm diam. Alatae (BL 0.8-1.0 mm, with
orange abdomen) differ from Ph. devastatrix in characters of the wings and
antennae (see Pergande, 1904).

Phylloxera glabra (von Heyden), Plate 2a Monoecious holocyclic, mainly

on Quercus robur. Fundatrices feed in spring from main veins, sometimes
causing inward curling of leaf-edge as in Ph. confusa. Subsequent generations
include yellowish-orange apterae (BL 0.7-0.85 mm) - which lay circular
clusters of eggs on the underside of the leaf - and both apterous and (later)
alate sexuparae (Grassi, 1912). Feeding causes necrotic spots on leaves. In
Europe (England, France, Germany, Italy) and introduced into New Zealand,
where it is recorded from Q. dentata (Sunde, 1984). It is regularly preyed upon
by Pullus auritus and Conwentzia psociformis.
Phylloxera globosa (Shimer) Galls on leaves of Carya amara (= cordiformis)
in Illinois were mostly rather large, globular and projecting equally on both
sides of leaf, or conical and projecting only below leaf, with flat circle above
('form conifera'); smooth, leathery, pale yellowish-green to dark green, with
a small central opening below leaf surrounded by small bracts. [According
to Pergande (1904), the smaller galls included in Shimer's (1867) original
description were those of Ph. caryaesemen.] Alatae (BL c. 0.6mm, with
orange-yellow abdomen), emerge very late in year, in late September-October.
Life cycle unknown. 2n = 6 (Steffan, 1968).

Phylloxera ilicis Grassi, Plate le, f Anholocyclic on Quercus ilex in central

Italy. Several generations of yellow apterae occur through spring and summer
feeding mainly on current and one-year-old twigs and causing small tuberosi-
ties; but sometimes also on foliage causing necrotic spots like Ph. quercus.
Alate virginoparae occur in autumn generations (Grassi, 1912).

Phylloxera intermedia Pergande Galls on leaves of Carya alba (= ovata) are

circular, reddish or yellowish, 3-6mm in diam., convex to conical on upper
side of leaf without any trace of a central nipple, paler and almost flat with
a small closed nipple on underside. Alatae (BL 0.8-1.3 mm, with pale orange
abdomen) emerge in late May-June, mainly from a jagged opening formed
by splitting of the upper surface of the gall (Pergande, 1904). Locality pro-
bably in vicinity of Washington, DC. Life cycle unknown.

Phylloxera italica Grassi Monoecious holocyclic on Quercus robur and

Q. petraea, only known from central Italy. Fundatrices develop in April-May
and the next two generations consist only of apterae (whitish, BL c. 1.5 mm),
which feed on undersides of leaves causing small, sparse, whitish spots.
Sexuparae are mostly apterous (Grassi, 1912).

Phylloxera kunugi Shinji Apterae (fundatrices?) are bright yellow, BL 2.3-

2.5 mm. They feed in spring from mid-rib on undersides of young leaves of
 Phylloxera 829

Quercus spp. (acutissima, variabilis) in Japan, causing the infested leaves to

curl downward. All their progeny are alate (BL 1.8-2.5 mm), and apparently
migrate to another host; the full life cycle remains to be established (Miyazaki
and Teramoto, 1991).

Phylloxera minima (Shimer) Galls on Carya amara (= cordiformis) in

Illinois, USA were small, round (diam. 1.1-5.0 mm) and numerous, with part
protruding above leaf shining brightly and a small opening beneath. Alatae,
present in the galls in early July, were translucent, yellowish-white with minute
dark spots on anterior part of body (Shimer, 1869). Life cycle unknown.

Phylloxera notabilis Pergande Galls on leaves of Carya illinoensis are ovoid

to spherical, bladder-like, very variable in size (diam. 2-18 mm), situated close
to mid-rib, projecting on both sides of leaf, green on top with a central nipple,
and reddish-green beneath covered with dense short yellowish hairs. Alatae
(BL 0.6-0.8 mm, with dusky wings and yellow-orange abdomen), emerge from
the top of the gall, which generally splits into long slender bract-like segments
in June-July. Monoecious holocyclic, with several generations of galls each
year; the smaller, secondary galls are produced mainly on seedling trees
(Stoetzel and Tedders, 1981). Brachypterous and migratory alatae, sexuparae
and sexuales are all produced in the primary galls. Sexuales lay overwintering
eggs on bark (Whitehead and Eastep, 1937). In eastern and southern USA,
and also recorded on C. illinoensis in southeast Europe (Georgia: Dzhibladze,
1975, as Xerophylla notabilis). Andersen and Mizeli (1987) studied the phy-
siological effects of galling on the host.

Phylloxera perniciosa Pergande Galls on young twigs, leaf petioles and

along mid-ribs of leaflets of Carya tomentosa; elliptical, with a blunt tooth-
like or thorn-like projection. In Maryland, Virginia, District of Columbia
and Pennsylvania; large gall populations in Virginia caused defoliation of
trees (Pergande, 1904). Also reported from C. illinoensis in Louisiana and
Mississippi (Carpenter et al., 1979), although the identity of these populations
with Pergande's species needs confirmation. Monoecious holocyclic, with
three generations per year; fundatrices, alate sexuparae and sexuales. Alate
sexuparae emerging from galls in May-early June laid eggs indiscriminately
on trees and surrounding vegetation. Sexuales laid overwintering eggs in
June, mostly high in trees on twigs that had been previously galled. The eggs
were preyed upon through the summer by various natural enemies, including
mites of six different genera. Surviving eggs hatched the following April
(Pergande, 1904).

Phylloxera picta Pergande Galls on leaflets of Carya tomentosa resemble

those of Ph. intermedia, but have thinner walls, and a central nipple above
as well as below the leaf; colour variable, diam. 3-6 mm. Alatae (BL 0.8-
1.3mm, with pale orange abdomen) emerge from an opening on the upper
side in late May-June (Pergande, 1904). Locality not stated; probably in
vicinity of Washington, DC. Life cycle unknown.
830 Phylloxera

Phylloxera pilosula Pergande Galls on leaflets of Carya glabra near

Washington, DC were flat or with a central depression above leaf and pro-
jecting with a well-developed central nipple below leaf; diam. 3-6 mm, pale
green to whitish, both sides densely covered with long, soft, glistening,
yellowish-white hairs. Alatae (BL c. 1.2mm, with orange abdomen) emerge
from the underside of the gall in early June (Pergande, 1904). Life cycle
unknown.

Phylloxera querceti Pergande Apterae (= fundatrices?) yellow to orange,

BL c. 0.5 mm. On undersides of leaves of Quercus spp. (alba, dentata(?),
macrocarpa) in the vicinity of Washington, DC (Pergande, 1904; Duncan,
1922). Affected leaves speckled with small yellow spots, especially near mid-
ribs and along smaller veins. Alatae of BL 0.6-1.0 mm, with orange abdomen.
Life cycle unknown.

Phylloxera quercina Ferrari (= spinulosa Targiani-Tozzetti) Apterae pale

yellow, with dorsal processes whitish (but darker in spring and autumn genera-
tions); BL 0.9-1.5 mm. On undersides of leaves of Quercus spp. (cerris,
ithaburensis, suber) in the Mediterranean region (Italy, Israel, Portugal),
causing yellow spots. Monoecious holocyclic; on Q. cerris in central Italy,
three-six parthenogenetic generations occur annually with the proportion
of alate sexuparae increasing in successive generations (Grassi, 1912, as
Acanthaphis spinulosa).

Phylloxera quercus Boyer de Fonscolombe (= florentina Targiani-Tozzetti)

Apterae greenish-yellow to orange-yellow; BL 0.7-0.9 mm. On undersides of
leaves of Quercus spp., causing yellowish spots. In southern Europe, North
Africa and southwest Asia. The life cycle is complex (Grassi, 1912). Over-
wintering usually occurs as eggs on evergreen oaks, Q. coccifera or Q. ilex.
Progeny of the fundatrices include alatae which fly in May to colonize
deciduous oaks for the summer months (petraea, pubescens, robur), and
also apterae which continue generations on Q. ilex, etc. In both cases alate
sexuparae appear in the fourth and fifth generations, and those produced on
deciduous oaks mostly return to evergreens to give rise to sexual morphs.
Overwintering as young larvae may also occur (see Grassi, 1912 for further
details). Mohammed and Abdulla (1985) studied the ecology of populations
identified as Ph. quercus in Iraq.

Phylloxera reticulata Duncan Apterae bright orange-yellow, with reddish

eyes; BL c. 0.8 mm. On undersides of leaves of Quercus kelloggii in California,
USA, dispersed along veins. Only apterae described (Duncan, 1922) and not
recorded since.

Phylloxera rileyi Riley Apterae dark brown with dorsal tubercles almost
black (but paler forms may occur); BL c. 0.6 mm. On undersides of leaves of
Quercus alba and Q. obtusiloba (= stellata) in Missouri, causing white or
yellow circular spots. Apparently monoecious holocyclic; alatae were pro-
 Phylloxera 831

duced from late July to October, and those of later generations are presumably
sexuparae, because sexuales are recorded laying overwintering eggs on twigs
(Riley, 1875; Pergande, 1904; Duncan, 1922).

Phylloxera rimosalis Pergande Galls on Carya tomentosa in eastern USA

(Washington, DC) are fairly large and conspicuous (diam. 3-11mm), pale
green sometimes tinged with red, usually on terminal leaves of young shoots;
flat to slightly convex above leaf, more prominently convex with a central
nipple beneath. Alatae (BL c. 1 mm, with pale orange abdomen) emerge in late
May-June through a round opening on the underside (Pergande, 1904). Life
cycle unknown.

Phylloxera russellae Stoetzel Galls on Carya illinoensis in southern USA are

circular, flattened, diam. 3.2-7.2 mm, with a reticulate surface pattern, pro-
jecting from both sides of leaflet with an opening on the underside marked
by white pubescence (Stoetzel, 1993). Galls mature and break open in May.
No alatae are known. Monoecious holocyclic; apterous sexuparae produce
sexuales which develop in the gall in May and produce overwintering eggs in
galls, on twigs, etc. (Stoetzel, 1981). Heavy populations of galls can cause
defoliation. Calcote (1983) studied resistance of pecan clones to Ph. russellae.

Phylloxera similans Duncan Apterae of BL c. 0.7 mm, appearance in life

unrecorded. Collected on Quercus macrocarpa in New York State (Duncan,
1922) and not recorded since original description. Biology unknown.

Phylloxera spinuloida Pergande Galls on petioles or at bases of main veins

of leaflets, of an undetermined Carya species in Florida, were globular (diam.
5-15 mm), with a dense covering of long fleshy filamentous outgrowths;
usually encircling the petiole and causing it to bend or curve. Colour uniformly
greenish-yellow, or brownish-yellow on one side. Galls opened in late March,
releasing alatae (BL 0.6-1.8 mm, with brownish wings and orange abdomen).
Life cycle unknown, although the early departure of alatae suggests host
alternation.

Phylloxera stanfordiana Ferris Apterae pale yellow, BL c. 0.75 mm. On

twigs of Quercus douglasii in California in September 'concealed beneath any
chance covering, especially numerous under the twig-encircling egg masses of
a moth' (Ferris, 1919). Other morphs and life cycle unknown.

Phylloxera stellata Duncan Apterae (BL c. 0.5mm, colour unrecorded)

caused brown spots on undersides of leaves of a Quercus sp., margaretta
(= stellata or alba) in mid-August in Texas, USA (Duncan, 1922). Other
morphs and life cycle unknown.

Phylloxera symmetrica Pergande Galls on leaves of Carya tomentosa and

other Carya spp. are variable; typical form is circular and almost flat above
leaf, protruding as a thick-walled pale cone below leaf, basal diam. 2-5 mm.
832 Phylloxerina

Alatae (BL 0.9-1.2 mm, with orange abdomen) emerge in late May-early June
through a small opening in the apex of the cone, after which the galls become
dry and brown. In vicinity of Washington, DC, USA. Life cycle unknown.
Pergande (1904) described variants from the typical form of gall ('vasculosa'
and 'purpurea') differing in colour, size and shape, which he considered to
be due to differences in hatching time of the fundatrices and developmental
stage of the leaf at onset of feeding.

Phylloxera texana Stoetzel Galls on leaves of Carya illinoensis are globular,

projecting mainly on upper side of leaf, with a small point below; diam. 4-
6mm (Stoetzel, 1981). Alatae (BL 1.0-1.6 mm, with golden-yellow abdomen)
emerge in mid- to late May through a large opening in the top of the gall,
which splits into bracts. In Texas, USA. Heteroecious holocyclic; alatae from
galls colonized undersides of leaves of Quercus spp. (falcata, virginiana), these
colonies producing both alatae and apterae in June (Stoetzel, 1985b). The
return migration to pecan has not been observed.

Phylloxera tuberculifera Duncan Apterae (BL c. 0.5 mm, colour unrecorded)

caused brown spots on undersides of leaves of Quercus havardii in Texas,
USA, in August (Duncan, 1922). Other morphs and life cycle unknown.

PHYLLOXERINA Börner Phylloxeridae

About seven species on Salicaceae, mostly living in dense white wax. The
species on Salix in Germany were studied by Iglisch (1965).

Phylloxerina capreae Börner, Plate 2b Apterae yellow to brownish-yellow,

under dense coat of white wax filaments; BL 0.6-0.9 mm. On Salix caprea
stems and branches, in bark crevices or under scales. Only known from
Germany (although a phylloxerid on Salix lanata in England, BMNH colln,
leg. VFE, has been provisionally assigned to this species). Monoecious
holocyclic (see Iglisch, 1965 for life cycle details).

Phylloxerina daphnoidis Iglisch Apterae light brown, under dense coat of

white wax filaments; BL 0.6-0.9 mm. On Salix daphnoides and S. elaeagnos
in Germany, living in bark crevices, etc. Monoecious holocyclic; Iglisch (1965)
gave life cycle details and compared Ph. daphnoidis with the very similar
Ph. capreae. The two species cannot be reliably distinguished on morphology
alone.

Phylloxerina nyssae (Pergande) Galls are usually crescent-shaped, yellowish-

white, formed in spring by rolling, scalloping and swelling of edges of leaves
of Nyssa sylvatica (Stoetzel, 1993). Later in the year (September), lemon-
 Phylloxerina 833

yellow apterae, BL 1.0-1.3 mm, completely covered in white wax wool, occur
together with sexuales in bark crevices of young trees, particularly in cavities
caused by dropping out of dead twigs about 2 m above ground level. Monoe-
cious holocyclic on N. sylvatica. In Maryland and District of Columbia, USA
(Pergande, 1904).

Phylloxerina popularia (Pergande) Apterae are pale greenish-yellow, covered

with white cottony wax, BL c. 1.4mm, living in deserted galls of Pemphi-
gus populitransversus and Pemphigus populicaulis, in bark crevices or on
branches and twigs, of North American cottonwoods (Populus monilifera),
P. fremontii and P. trichocarpa) in Louisiana, Michigan, Missouri, South
Dakota, Texas and California (Pergande, 1904; Davidson, 1915). Monoecious
holocyclic; apterous sexuparae and sexuales were found in old Pemphigus
galls in October (Pergande, 1904).

Phylloxerina populi (Del Guercio) Apterae yellow, covered in wax wool.

On bark of trunk and branches of Populus alba, only known from Italy.
Monoecious holocyclic. Bonfigli (1909; as Pseudochermes populi) compared
it with Ph. salicis.

Phylloxerina prolifera (Oestlund) Apterae very pale lemon yellow, found

in autumn in empty galls of Pemphigus populicaulis in Minnesota, USA
(Oestlund, 1887). Not recognized since original description and possibly
not distinct from Ph. popularia, despite the apparent differences noted by
Pergande (1904).

Phylloxerina salicicola (Pergande) Apterae yellow to brownish-yellow,

covered with white wax wool; BL 0.6-0.9 mm. Originally described from bark
crevices or other cracks in stems and twigs and sometimes also on undersides
of leaves, of an unidentified Salix species (either discolor or humilis) in eastern
USA (Pergande, 1904). Monoecious holocyclic, with sexuales in September.
Davidson (1915) recorded this species from Salix lasiolepis in California,
usually living in bud axils on twigs but also on one occasion heavily infesting
bared roots on a stream bank. These Californian populations appeared to be
anholocyclic and could possibly have been introduced on European Ph. salicis.

Phylloxerina salicis (Lichtenstein) Apterae pale green, yellow or brownish,

secreting profuse white wax wool; BL 0.7-1.2 mm. Living in bark crevices on
trunk and branches of Salix alba in Western, southern and Central Europe,
and across temperate Asia to China (Zhang et al, 1985) and eastern Russia
(Pashchenko, 1988b). Introduced to Victoria, Australia (Eastop, 1966) and
perhaps to North America (Börner and Heinze, 1957). Monoecious holocyclic
in Europe; sexuales in September (Iglisch, 1965).

Phylloxerina sp. Holman (1974) described (but did not name) a species of
Phylloxerina on trunks of Salix longipes in Cuba. Life cycle unknown. Similar
to European Ph. capreae (but not compared with the North American species).
834 Pineus

PINEUS Shimer Adelgidae

Clearly differentiated from the other adelgid group, Adelges, by their biology -
e.g. gall structure, secondary host association with Pinus - and by various
morphological features. The most consistent character that applies to all
morphs is the reduction in the number of abdominal spiracles to only four
evident pairs. The galls, usually on shoot tips of Picea, are generally less
compact than those of Adelges and have intercommunicating chambers. Adult
apterous morphs on secondary hosts (sistens and progrediens) are very broadly
pear-shaped - often almost globular - and have a fused and pigmented head
and prothoracic shield.
Pineus species may be holocyclic and host-alternating between Picea and
Pinus, completing a cycle with seven morphs in two years as in holocyclic
Adelges; or they may be anholocyclic on either Picea or Pinus (except for
Pineus abietinus, which is anholocyclic on Abies). There is also one species,
Pineus floccus, that is now known to have an anholocyclic, annual host
alternation between Picea and Pinus. Knowledge of the life cycle of many
species is still incomplete. Several anholocyclic species on Pinus produce alate
sexuparae which appear to be non-functional, either because they are unable
to produce males or because they fail to find a suitable Picea species to act
as primary host.
Systematic accounts of Pineus are available for Britain (Carter, 1971),
Europe (Börner and Heinze, 1957; Heinze, 1962), India (A.K. Ghosh, 1984a),
Japan (Inouye, 1953) and North America (Annand, 1928).

Pineus abietinus Underwood and Balch Only the first instar and adult
progrediens (aestivoprogrediens) are described; adults are very small (about
0.6mm), light reddish-brown with dark cephaloprothoracic shield, secreting
white wax wool, in bark crevices on stems and branches of Abies amabilis and
A. grandis in the northwest coastal region of North America (Washington,
southern British Columbia). This is the only known Pineus associated with
Abies. Wax pores are circular and discontiguous. Apparently anholocyclic and
monomorphic, probably with two or three progrediens generations per year
(Underwood and Balch, 1964).

Pineus boerneri Annand Apterae very small, reddish-brown, laying eggs on

pine twigs in abundant greyish-white wax wool. Described from Pinus radiata
in California (Annand, 1928), but possibly East Asian in origin, where it has
mainly been recorded under the name P. laevis (see McClure, 1984b); also
now in northeastern USA, Hawaii, South America (as Pineus havrylenkoi
Blanchard), Australia, New Zealand and more recently East Africa (as
Pineus pini). Pineus simmondsi Yaseen and Ghani, on Pinus roxburghii in
Pakistan (Yaseen and Ghani, 1971), could also be this species. Anholocyclic
 Pineus 835

everywhere; introduced populations attack a range of exotic pines from both

Europe (e.g. P. halepensis) and North America (e.g. P. radiata, P. caribaea).
In Australia and New Zealand the name laevis has been applied to both
boerneri, and pini (q.v.), which are difficult to differentiate by morphology
alone. Biological control of boerneri and/or pini has been successful using
Leucopis sp. (Chamaemyidae) in New Zealand, Chile and Hawaii (Mills,
1990). Dynamics of East African populations were studied by Mailu et al.
(1980; as pini), and in the rather different conditions of Pinus resinosa
plantations in Connecticut, USA, by McClure (1989b, 1990). 2n = 16*
(California, Hawaii) or 17* (California, Africa, Australia, New Zealand).

Pineus boycei Annand This species forms 2-4 cm long galls on Picea
engelmanni, green with a reddish tint at the bases of the needles, usually at
the shoot tips. It is only recorded from Montana and Oregon, USA. Alate
gallicolae emerging from the galls in the second half of July are 1.7-2.5 mm,
with abundant wax posteriorly on the reddish-brown abdomen (Annand,
1928). Their destination is unknown. No other morphs or life cycle details are
known (although it seems possible that alate specimens described by Annand
under the name Pineus engelmanni are the sexuparae of this species).

Pineus cembrae (Cholodkovsky) (= sibiricus Cholodkovsky) Host-alter-

nating between Picea spp. (abies, glehnii, obovata) and stone pines, Pinus
spp. of subsection Cembrae (cembra, pumila, koraiensis, sibirica); in alpine,
Central and Eastern Europe, and across Russia to Japan. Other pines are
not usually attacked (Potapova, 1984), except that P. parviflora is recorded
as a host in Japan. Galls (Fig. 132A) are 4-5 cm long, often asymmetrical,
distorting shoot tips, with needles concentrated on one side, somewhat
resembling the galls of Adelges cooleyi. Gallicolae are dark red and secrete
abundant wax, migrating to Pinus in June in Germany (Börner and Heinze,
1957), but in August in Japan (Inouye, 1953). The neosistentes overwinter
mainly near ends of twigs at the bases of last year's pine needles, giving rise
the next year to alate sexuparae and one or two generations of apterous pro-
gredientes living in dense wax wool on new shoots. Inouye (1953) described
and illustrated all morphs. A form collected on the needles of P. koraiensis
in Heilongjiang Prov., China is described as a subspecies, P. cembrae pini-
koreanus (Zhang and Fang, 1981). 2n = 18*.

Pineus cladogenous Fang and Sun Anholocyclic on Pinus koraiensis in

Heilongjiang Prov., China, overwintering in fissures in the bark. The adult
hiemosistentes (collected in April) are small and ladybird-shaped; BL about
0.6mm (Fang and Sun, 1985). Perhaps closely related to P. harukawai.

Pineus coloradensis (Gillette) Anholocyclic on numerous Pinus spp. in USA

and Canada. Apterous progredientes feeding on the needles secrete abundant
wax, which in summer becomes covered with a dark mould. One hiemopro-
grediens and two aestivoprogrediens generations occur in New England, where
this adelgid has been injurious to plantations of P. resinosa (McClure, 1989b).
836 Pineus

Fig. 132. Galls of Pineus on Picea. A, Pineus cembrae on Picea abies (after Francke-Grosmann, 1938);
B, P. orientalis on Picea orientalis (after Carter, 1976); C, P. pinifoliae on Picea mariana (after Patch,
1910a; open gall on left, closed gall on right).

Alate sexuparae, presumed non-functional, are recorded from Colorado, but

not from California or Connecticut. Descriptions of morphology and biology
in comparison with related species are given by Annand (1928), Doane (1961)
and McClure (1989b). McClure (1984a) studied interactions between this
adelgid and Pineus boerneri on Pinus resinosa, and Huff and McDonald
(1977) studied differences in its susceptibility to 19 white pine species.

Pineus cortecicolus Fang and Sun Anholocyclic on Pinus koraiensis in

Heilongjiang Prov., China, overwintering under bark in only a little wax
(Fang and Sun, 1985). BL of adult hiemosistens about 0.85mm. From the
description this species appears similar to P. hosayai, but differs in over-
wintering site and amount of wax.

Pineus floccus (Patch) 'Host-alternating' between Picea spp. (rubens,

mariana) and Pinus strobus in eastern USA, but the host alternation revealed
by life history studies in Virginia is remarkable because it occurs annually and
only involves parthenogenetic morphs; there are no fundatrices, sexuparae or
sexuales (Walton, 1980). There is a single generation (gallicolae) on Picea, and
a series of progrediens generations on pine. Overwintering apterous progre-
dientes on P. strobus produce a spring generation of both apterous and alate
progredientes, and the latter fly to spruce buds in May. Galls are induced
by the developing gallicolae. They are usually terminal, 1.8-4.5 cm long,
resembling those of Adelges cooleyi but shorter, stouter and looser; green, but
acquiring a purplish or reddish tinge as they mature, the protruding ends
of the needles remaining green and of normal length. Adult gallicolae, BL
0.9-2.0 mm with abundant wax on head and thorax and a yellow-brown
abdomen, migrate in July to both old and new needles of Pinus strobus.
Apterous adults of the second generation on pine overwinter at the bases of
buds, needles or stem scales, moving to growing points in spring. In New
York, however, where heavy galling of Picea rubens in plantations is recorded,
 Pineus 837

Deboo and Weidhaas (1967) believed that gallicolae were recolonizing spruce
directly and not migrating to pine.

Pineus ghanii Yaseen and Ghani Anholocyclic on Pinus griffithii in the

Galis and Kaghan valleys, Pakistan, attacking stems and twigs of mature and
old trees (Yaseen and Ghani, 1971). There are two generations per year, a
hiemosistens becoming adult in April, and an aestivosistens becoming adult
in August-September. Adults are light reddish-brown, BL 0.9-1.2 mm.

Pineus harukawai Inouye Anholocyclic on white pines (Pinus parviflora,

P. strobus) in Japan (Hokkaido, Honshu, Kyushu), on stems or older
branches of young pines, or at the bases of the needles on young branches
of older pines. Only the adult of the overwintering sistens - about 0.9 mm and
orange to brownish-red with abundant wax, laying large numbers of eggs in
April to May - is described; other details of the life cycle are unknown. This
species is sometimes injurious in pine nurseries in Hokkaido (Inouye, 1953).

Pineus hosoyai Inouye Only the apterous progrediens and immature sex-
upara are described, collected from Pinus koraiensis in June in Korea and
from Pinus sp. in May in China (Inouye, 1953). The overwintering sistentes
were found in February in Korea, living in abundant wax on shoots of the
previous year's growth. Life cycle is unknown.

Pineus matsumurai Inouye Apterous progredientes and alate sexuparae

develop in June at needle bases on young branches of Pinus thunbergii and
P. densiflora in Japan (Inouye, 1953). The apterous progrediens is small
(about 0.8 mm), dark reddish-brown, covered in dense fine wax wool. The
destination of the sexuparae and the rest of the life cycle are unknown. This
species seems to be close to Pineus strobi; differences are discussed by Inouye
(1953).

Pineus orientalis (Dreyfus), Plate 1d Host-alternating between certain Picea

spp. (orientalis, polita, jezoensis, but not abies) and various Pinus spp. of the
subsection Sylvestres (densiflora, mugo, sylvestris, thunbergii) from Western
Europe across Asia to Japan. Galls (Fig. 132B) are terminal, shorter and
usually straighter than those of P. cembrae, with protruding needle tips more
evenly distributed around the gall. Gallicolae fly to Pinus in late June to
August and lay eggs on the previous year's needles, the eggs often being
covered by the wings of the dead alata (C.I. Carter, pers. comm.). There are
numerous accounts of the biology of this adelgid, which is not differentiated
from its anholocyclic close relative P. pini in some of the earlier literature
(e.g. in Europe: Börner, 1908; Marchal, 1913; van der Goot, 1915; Mordvilko,
1935; and in Japan: Inouye, 1953). 2n = 20*.

Pineus pineoides (Cholodkovsky) Anholocyclic, with one summer and one

winter generation per year, both of apterae which live scattered on well-shaded
parts of the bark of the trunk and larger branches of Picea spp. (abies,
838 Pineus

obovata, omorika, rubens). The insects are yellowish-grey or dark red-brown

to almost black, dorsoventrally flattened, with the cephaloprothoracic shield
of strongly pigmented specimens heavily sclerotized; BL 0.6-0.8 mm. Each
lays its eggs under a separate tuft of wax wool. The overwintering genera-
tion becomes adult in May; the adults of the summer generation, similar
except that they have rather fewer wax pores, may be found in July. In
Europe, Canada and also recorded from bonsai spruce in Australia (New
South Wales; BMNH colln, leg. D. Koreshoff). For details of biology see
Steffan (1963, in Germany) and Underwood (1963, in Canada). Pineus
konowashiyai, on Picea abies in Japan, has a similar habitat and life cycle to
P. pineoides, but differs in having large discontiguous wax pores (Inouye,
1953). 2n = 22 (Steffan, 1968) or 17*.

Pineus pini (Macquart) (= laevis Maskell), Plate lc Anholocyclic on Pinus

sylvestris and Pinus mugo in Western and Central Europe, with an over-
wintering (hiemosistens) generation on the twigs and two or more overlapping
generations of progredientes attacking the current year's shoots. Eggs are laid
in abundant wax wool. The second progrediens generation in May-June
includes alatae, but first instar crawlers dispersed by wind may be a more
common means of infection of young seedling pines (C.I. Carter, pers.
comm.).
The morphs of P. pini are all virtually indistiguishable from the equivalent
secondary host generations of P. orientalis, and it seems probable that the
anholocyclic P. pini arose from the holocyclic P. orientalis by loss of a func-
tional sexual generation, as suggested by Marchal (1913). Pineus pini in
Europe seems to restrict its attacks mainly to Pinus sylvestris and Pinus mugo,
but forms with a very similar morphology and biology are found in North
America, Australia and New Zealand, and can be injurious to a wide range
of Pinus species. In the Australian literature populations of P. pini group
(= laevis) are confused with P. boerneri, which also occurs on a range of
pines in both Australia and New Zealand. Maskell (1885) clearly thought that
the species he described as laevis was the pest on Pinus radiata, etc., but it
seems that the specimens he described were P. pini, both species having been
introduced to New Zealand. 2n = 22 (Steffan, 1968, or 20* (in Europe), 21(?)*
in Australia and 19* in New Zealand.

Pineus pinifoliae (Fitch) Host-alternating between Picea spp. (engelmanni,

glauca, mariana, pungens, rubens, sitchensis) and white pines (Pinus strobus,
P. monticola) in North America. The galls on spruce (Fig. 132C) are conical,
1.8-4.3 cm long, with their constituent needles characteristically flattened and
broadened to form 'scales'. They are purple and green at first, becoming
reddish-brown at the time of opening in mid-June to early July, when the
needle scales flare outwards. The gallicolae, BL 1.6-2.3 mm, fly to needles of
the previous year's growth of white pine; they invariably face the base of the
needles and deposit their eggs beneath their wings, dying after oviposition with
the wings still protecting the eggs. The first instar sistentes (neosistentes) move
to the axes of new shoots to overwinter. There are two apterous progrediens
 Pineus 839

generations on pine, alate sexuparae being produced in May to fly to old

needles of spruce. For further life cycle details see Balch and Underwood
(1950). Dynamics of the gall populations on spruce was studied by Howse and
Dimond (1965) and of the populations on white pine by Dimond and Alien
(1974). The histology of feeding damage to white pine shoots has also been
studied (Alien and Dimond, 1968). 2n = 22 according to Steffan (1968).

Pineus sichuananus Zhang Described from Picea likiangensis and Picea

purpurea in Sichuan, China. The first instar and adult sistens, which one
would normally expect to occur on a Pinus species, were recorded along with
galls and gallicolae on spruce, so the life cycle needs clarification. Pineus
sichuananus is considered to be closely related to P. cembrae; the differences
given all involve the wax glands (Zhang, Zhong and Tian, 1980).

Pineus similis (Gillette) Anholocyclic, living without host alternation on

a wide range of Picea spp. in North America (abies, glauca, engelmanni,
mariana, pungens, rubens) and introduced into Britain where it also colonizes
Picea sitchensis (Carter, 1975). Cumming (1962a) studied the unusual life cycle
of this adelgid in Canada. Pseudofundatrices (not true fundatrices, because
they hatch from unfertilized eggs) overwinter in second or third instar under
bud scales. Their feeding in spring causes rather loose galls 1-4 cm long, with
the constituent needles much paler than the normal new growth. As the gall
matures the shoot and needles become brown and twisted giving it a ragged
appearance. Uniquely, both alate and apterous gallicolae are produced in the
galls. Apterous gallicolae oviposit within the galls, but apterae may also be
found outside the galls, laying eggs in small tufts of wax on the stem. Alate
gallicolae, BL 1.0-2.1 mm and dark reddish-brown, fly in June-July to
recolonize spruce needles, laying eggs in white wax wool which apparently
develop into the overwintering pseudofundatrix generation. 2n = 22 according
to Steffan (1968).

Pineus strobi (Hartig) Anholocyclic on Pinus strobus in North America and

introduced on this pine into Europe. In England, Carter (1971) has also
recorded it attacking one species of European white pine, Pinus peuce. Raske
and Hodson (1964) describe the life cycle and biology in Minnesota, USA and
their observations largely agree with those made in Europe (e.g. Marchal,
1913; Carter, 1971). Overlapping generations of sistentes and progredientes
feed in large aggregations through the spring and summer on shaded parts of
the bark of the trunk and undersides of branches, secreting copious white wax
wool. On seedling trees, parts of the new growth or last year's growth, where
the needles are densest, are colonized. Overwintering occurs as immature
apterae (progrediens or sistens) on the trunk, or under scales of dwarf shoots.
Alate sexuparae are often produced in May-June and fly to Picea mariana
to lay eggs on the needles. However, the sexual phase is abortive, either
because the sexuales hatch but fail to develop (Raske and Hodson, 1964) or
due to the absence of any males (Marchal, 1913). No galls or gallicolae of
P. strobi have been found on Picea mariana or any other spruce. Sluss and
840 Platyaphis

Foote (1973) studied predation by Leucopis pinicola (Chamaemyidae) and

other natural enemies of P. strobi in Ohio, USA. 2n = 22 (Steffan, 1968)
or 20*.

Pineus sylvestris Annand Apparently anholocyclic on Pinus spp. (radiata,

sylvestris) in California, USA (Annand, 1928). Inouye (1953) recorded what
appears to be the same species from Pinus thunbergii in Hokkaido, Japan,
but the differentiation from pini and boerneri is problematic. Reddish-brown
apterous progredientes, BL 1.0-1.1 mm, with rather sparse wax, were found
in May-June in California feeding inconspicuously at the needle bases near
male cones. Alate sexuparae appeared early in May, but their fate is unknown,
no generations on spruce having been identified. The overwintering stages on
Pinus, mainly second and third instar progredientes, were found under stem
scales and behind lateral buds near the ends of branches.

Pineus wallichianae Yaseen and Ghani Anholocyclic on Pinus griffithii

(= wallichiana) in the Galis and Swat Hills, Pakistan, attacking stems and
twigs of mature and old trees (Yaseen and Ghani, 1971). Adult sistentes are
light brown with a thick covering of wax, BL c. 0.9-1.0 mm. There are only
one or two sistens generations per year, and no alatae are known. Morphology
and life cycle are apparently very like Pineus ghanii, and this is possibly a
highland form of that species.

PLATYAPHIS Takahashi Drepanosiphinae(?): Phyllaphidini(?)

One species on Fagus in Japan. Only the very specialized apterous vivipara
is known, and its phylogenetic position is uncertain. Takahashi (1957) com-
pared it with Dasyaphis, but Shaposhnikov and Gabrid (1987) suggested that
it belongs in the Hormaphidinae.

Platyaphis fagi Takahashi Apterae broadly ovate, flattened, green with pale
antennae and legs; BL 1.8-2.0 mm. On undersides of leaves, along mid-ribs,
on Fagus crenata in Japan (Takahashi, 1957). Life cycle and other morphs
unknown.

PLOCAMAPHIS Oestlund Aphidinae: Pterocommatini

A small genus of Salix-feeding aphids with a rather even holarctic distribution.

They are less hairy than Pterocomma and have flangeless siphunculi. Most
species secrete flocculent wax and they are not attended by ants. Richards
 Prociphilus 841

(1966b) revised the genus (which in his concept included Fullawaya), and Heie
(1986) reviewed the European species.

Plocamaphis amerinae (Hartig) Apterae yellowish, greenish or brownish,

wax-covered in life, with dark head, antennae and legs; BL 3.0-4.5 mm. On
young growth of Salix spp. (alba, purpurea, viminalis), sometimes causing
damage in osier plantations. In northwest and northern Europe, eastwards to
western Siberia. Monoecious holocyclic; apterous males and oviparae occur
in October-November. Populations with very short siphunculi and no dorsal
dark markings on a Salix species (probably lapponum) in Sweden were
described as a subspecies (P. amerinae borealis) by Ossiannilsson (1959); such
forms are also reported from S. viminalis in Poland (Szelegiewicz, 1965).

Plocamaphis coreana (Okamoto and Takahashi) Apterae dark yellowish-

green, covered with white wax; BL c. 3.5 mm. Described attacking branch and
stem of an unidentified Salix sp. in Korea (Okamoto and Takahashi, 1927;
Paik, 1965, as Clavigerus salijaponicus). Pashchenko (1988b) recorded it
from S. crassijulis and S. parallelinervis in eastern Russia (Kamschatica). Life
cycle unknown.

Plocamaphis flocculosa (Weed) Apterae grey or yellowish-grey, densely

covered with wax powder; BL 3.1-5.0 mm. Head, legs and antennae darker,
and siphunculi orange. On trunk and branches of Salix spp. in North America
(discolor, lasiolepis) and Europe (caprea, cinerea, nigricans, repens). Also
recorded from China. Three subspecies have been distinguished in the very
variable European populations (Ossiannilsson, 1959), but further work is
required to determine whether these are valid taxonomie categories (Heie,
1986). The European form known as P. flocculosa brachysiphon produces
oviparae and alate males in October (Stroyan, 1955; as P. goernitzi). Plo-
camaphis assetacea Zhang, described from Salix sp. in Tibet (Zhang and
Zhong, 1981b), is stated to be closely related.

Plocamaphis martini Richards Apterae light brown to pink (no wax men-
tioned); BL 2.8-3.5 mm. On roots of Salix sp(p). in arctic and subarctic
Canada (Baffin Is., Quebec). Monoecious holocyclic; apterous males and
oviparae occur in mid-August (Richards, 1963). Possibly conspecific with
P. amerinae.

PROCIPHILUS Koch Pemphiginae: Pemphigini

About 50 species ranging in body size from rather small to very large, usually
lacking siphunculi and with well-developed wax glands, even in fundatrices.
Their host plant associations are unusual, in that they have apparently trans-
ferred their sexual generation in relatively recent times from Populus to
842 Prociphilus

various more advanced 'primary' hosts, e.g. in Rosaceae, Caprifoliaceae and

Oleaceae, while mostly retaining conifer roots as secondary hosts for the
parthenogenetic generations. The species are fairly evenly distributed through
the northern hemisphere; about 20 in North America, 7 in Europe, 3 in Central
Asia and 10 in East Asia. The life cycles of many species are unknown.
Accounts are available for northwest Europe (Heie, 1980), India (A.K. Ghosh,
1984b), Japan (Inouye, 1956), China (Tao, 1970) and North America (Smith,
1974a). Our concept of Prociphilus includes Stagona and we also include as
subgenera Pulvius, Neoparacletus and Meliarhizophagus, each with a single
species, which all appear to be relatively recent biological deviates from the
main evolutionary lineage of Prociphilus.

Prociphilus americanus (Walker) Host-alternating between Fraxinus spp.

and roots of Abies (balsamea, procera) in North America. The type specimen
is an alate sexupara from Nova Scotia (Doncaster, 1961); prior to Doncaster's
redescription this aphid was called P. fraxinidipetalae and/or P. venafuscus.
Early records from roots of Pseudotsuga are probably in error (Smith, 1969)
and records from other trees (e.g. Aesculus) are probably of vagrant indi-
viduals. Fundatrices on ash are dark brown to black, globose, forming early
spring colonies in curled terminal leaves of new growth (Essig, 1911) or on
twigs (Patch, 1909); populations may also develop on suckers and at base of
trunk 'at quite a depth underground' (Essig, 1911). Emigrant alatae maturing
from late April to early June are very dark green to black with dusky, bluish-
tinged wings and long bluish-white wax flocks extending posteriorly; BL
2.8-4.8 mm. They found wax-covered colonies on Abies roots. Sexuparae
return to produce dwarf sexual morphs on trunk of Fraxinus in late August-
October (Smith, 1969). Patch (1909) described the sexual morphs (as vena-
fuscus) and recorded that they were also deposited in large numbers on
Syringa, which may be an alternative primary host.

Prociphilus bumeliae (Schrank) Host-alternating between Oleaceae (usually

Fraxinus excelsior, sometimes Ligustrum vulgare, Syringa vulgaris) and roots
of Abies. Fundatrices are brown, covered with white wax wool, forming loose
'leaf nests' on primary host in spring. Spring colonies also form on bark of
stems and branches. All progeny alate, with blackish-brown head and thorax
spotted with white wax, and light brown to yellowish-red abdomen (Heie,
1980); BL 3.8-5.5 mm. They migrate in May-June to found wax-covered
colonies on Abies roots. Sexuparae return to Fraxinus in late August-
September (Mordvilko, 1935). Widely distributed in northwest, Central and
Eastern Europe. Records from East Asia should probably all be referred to
P. oriens. [Alatae of Prociphilus 'sp. B' on Fraxinus floribunda in northern
India (A.K. Ghosh et al., 1970) key to bumeliae, except that they lack dis-
cernible dorsal cephalic wax glands and have many more secondary rhinaria
(45-48) on ANT III.]

Prociphilus carolinensis Smith Apterae oval, presumably wax-covered in

life, with protrudiag dark CAUDA and anal plate; BL c. 2.5 mm. On roots of
 Prociphilus 843

Pinus spp. (including P. taeda), 3-4 feet below ground, attended by ants.
Sexuparae (BL 2.8-3.3 mm) emerge in October-December, migrating to an
unknown primary host. Sexuales are deposited and eggs laid on the bark of
various trees, especially Malus, but apparently do not hatch (Smith, 1969). In
USA (North Carolina) and Canada (Manitoba).

Prociphilus caryae caryae (Fitch) (= alnifoliae Williams) Spring popula-

tions (fundatrices + immature alatae) cup or curl the leaves of Amelanchier
spp. and turn them yellow or whitish. (Records from Crataegus are pro-
bably due to misidentification, particularly confusion with P. corrugatans.)
Emigrant alatae (BL 2.7-3.3 mm) fly in June to colonize roots of Pinus
sp(p)., sexuparae returning to Amelanchier in September-October. In western
USA and Canada (British Columbia). (No field observations have been made
of these western populations on pine roots, but Smith (1969) obtained colonies
readily on Pinus strobus in the laboratory, and sexuparae collected on
P. monticola in British Columbia (BMNH colln, leg. G. Shrimpton) appear
to be this species.) Similar aphids in eastern USA are regarded as a subspecies.

Prociphilus caryae fitchii Baker and Davidson Spring populations

(fundatrix + immature alatae) cause cupping and curling of leaves of Malus
spp. (pumila, angustifolia) and Amelanchier spp. Emigrant alatae (BL 1.8-3.3
mm) migrate in June to found ant-attended colonies on roots of Pinus strobus.
Sexuparae return to the primary hosts in October-November where they
deposit sexuales in bark crevices near base of tree or in nearby leaf litter
(Smith, 1969). Parthenogenetic populations also persist throughout the year
on white pine roots. A third closely-related form, P. caryae arbutifoliae, forms
spring populations on Photinia arbutifolia in California, from which alatae
(BL 2.9-4.1 mm) migrate to an unknown secondary host, probably pine roots.
[Records of P. caryae fitchii on Lonicera in India (A.K. Ghosh et al., 1981;
A.K. Ghosh, 1984b) apply to Prociphilus himalayensis.]

Prociphilus cornifoliae Singh, Das and Raychaudhuri Apterae (BL 4.2 mm,
colour in life unrecorded) were collected on roots of Cornus sp. in April in
Manipur, India (Singh et al., 1977; habitat information from A.K. Ghosh,
1984b). Alate specimens and immatures were also collected, both from the
same host, locality and date, and from an unidentified plant in Nepal. Life
cycle unknown.

Prociphilus (Neoparacletus) corrugatans (Sirrine) Spring populations cause

curling and cupping of leaves of Crataegus spp.; infested leaves turn pale
whitish, often with a reddish tinge at the edge. Fundatrices yellowish-green
with flocculent wax, with greenish immatures that develop into emigrant
alatae (BL 1.9-2.9 mm, with brownish abdomen). Migration occurs in May-
June to Juncus spp. (and to Carex?, where populations may persist through-
out the year. Sexuparae returning to Crataegus in autumn often deposit
sexuales on Malus, but apparently these do not produce spring populations
(Smith, 1972). Records from Amelanchier are probably all due to confusion
844 Prociphilus

with P. caryae. Widely distributed in North America. [Dark brown or reddish

curled leaves on Crataegus tomentosa, noted in the original description of
P. corrugatans (Sirrine, 1894), are considered by Smith (1972) to be due to
feeding by another Prociphilus sp., possibly the undescribed primary host
generations of P. erigeronensis.]

Prociphilus (Stagona?) crataegicola Shinji Described from Crataegus sp. in

June in Japan; alata has light greenish abdomen and antennae with secondary
rhinaria distributed III 22, IV 7, V 5 (Shinji, 1922a). Other morphs and life
cycle unknown, except that alatae collected from pine roots in Taiwan and
Japan by Tao (1970), as 'Prociphilus pini n. sp.' could be the sexuparae of
this species. Possibly crataegicola is a synonym of pini Burmeister; alatae
(BL 3.2mm) collected on 'apple' in Japan and identified by Takahashi as
crataegicola (BMNH colln, leg. T. Tanaka) have secondary rhinaria
distributed III 24-29, IV 6-8, V 6-8, and closely resemble European P. pini.
[P. sasakii Monzen, described from Malus pumila, was synonymized with
crataegicola by Shinji (1941), but this seems doubtful as Monzen's species had
siphunculi (unusual in Prociphilus) and less rhinaria on III.]

Prociphilus (Stagona) crataegistrobi Smith Fundatrices and their progeny in

spring curl leaves of Crataegus sp., which remain green but may tend to fade.
Progeny all alate, BL 2.2-2.5 mm, migrating in June to found colonies on
roots of Pinus strobus. The apterous exules live in individual wax-lined cells
associated with small roots and mycorrhizae (Smith, 1969). Sexuparae return
to trunks of Crataegus in October-December, but pine root-feeding popula-
tions also persist through winter. In USA; only known from North Carolina.

Prociphilus formosanus Takahashi In dense spring colonies distorting leaves

of Fraxinus sp. ('insularis') in Taiwan (Takahashi, 1935d). Alatae (collected
in May) have dark green abdomen, and BL c. 4 mm; antennae with secondary
rhinaria distributed III 32-39, IV 7-11, V 7-9, VI 0. Immatures yellow with
cottony wax on abdomen. Other morphs unknown. Alatae were also collected
on Picea morrisonicola, which is therefore thought to be the secondary host,
but confirmation of this seems necessary.

Prociphilus fraxini (Fabricius) Host-alternating between Fraxinus excelsior

and roots of Abies. Fundatrices feed at bases of new shoots of ash, giving
rise to large colonies of immature alatae which develop to maturity in dense
leaf nests, often high up in trees (Danielsson and Carter, 1993b). Alatae (BL
3.3-5.1 mm) fly in late June-July to found wax-covered colonies on Abies
roots, the apterous exules being almost indistinguishable from those of
P. bumeliae. Sexuparae return to Fraxinus in October-November. Widespread
in Europe; specimens from Turkey on Fraxinus oxycarpa (BMNH colln, leg.
H. Çanakçioglu) are possibly an undescribed, closely-related species.

Prociphilus (Meliarhizophagus) fraxinifolii (Riley) Monoecious on American

Fraxinus spp. Yellow-green to pale green fundatrices and their progeny (both
 Prociphilus 845

apterae and alatae) live in curled and clumped leaves at tips of twigs through-
out the growing season. Alatae (BL 1.4-2.8 mm, with yellow-green abdomen)
are produced in April-October. Ant-attended colonies also occur all year
round on Fraxinus roots. Alate sexuparae occur in October-November
(Smith, 1974a). Common and widely distributed in USA; also in Canada
(British Columbia, Manitoba), Mexico and introduced to Chile (Carrillo,
1977) and South Africa (on planted F. pennsylvanica and F. velutina; Müller
and Scholl, 1958, and BMNH colln). Carver (1980a) described an aphelinid
parasitoid on P. fraxinifolii in Iowa; Brundrett and Kendrick (1987) studied
symbiosis between root-feeding populations and a bolete fungus. 2n = 20
(Robinson and Chen, 1969) or 2n = 22*.

Prociphilus (?Neoparacletus) ghanii Hille Ris Lambers Apterae broadly

oval, appearance in life unknown; BL 1.3-1.7 mm. Collected on Salix
wallichiana in Pakistan (Hille Ris Lambers, 1973b). Other morphs, biology
and life cycle unknown.

Prociphilus (? Stagona) himalayensis Chakrabarti Described from sexuparae

(BL 2.6-3.4 mm, wax-covered) collected on roots of Pinus excelsa (= griffithii)
in Himachal Pradesh, India (Chakrabarti, 1976). It was placed in subgenus
Stagona, although the pterostigma is not elongate; perhaps because the
apterous exules described as paratypes (from an unidentified host) have a tibial
chaetotaxy like that of P. (5.) crataegistrobi. Banerjee and Chakrabarti (1993)
have confirmed experimentally the suspected (Chakrabarti et al., 1985) migra-
tion from folded-leaf galls on Lonicera quinquelocularis. The spring forms
had earlier been recorded as P. caryae fitchii by A.K. Ghosh (1984b). Samples
have been seen from the Punjab, Himachal Pradesh, Kashmir and Pakistan.
Rather similar spring migrants but with more rhinaria were collected from
Eriobotrya petiolata in West Bengal (BMNH colln, leg. A.N. Basu), and there
are apterae of other species on roots of Pinus griffithii in the Himalayas. The
group merits further investigation.

Prociphilus (Stagona) konoi Hori Host-alternating between Lonicera spp.

and Picea. Fundatrices yellowish-green, curling terminal leaves of Lonicera
longitudinally. All progeny are alate (BL 2.5-2.8 mm, with greenish-brown
abdomen) and migrate in June-July to found colonies on roots of Picea
glehnii (Hori, 1938; requires experimental confirmation). Sexuparae (with
secondary rhinaria distributed ANT III 15-16, IV 8-9, V 7-8, VI 3-4) occur
in October-November. In Japan and eastern Siberia (Pashchenko, 1988b).
2n = 18*.

Prociphilus kuwanai Monzen (= orientalis Mordvilko, 1929a) Feeding by

the fundatrix in spring on Pyrus spp. (and occasionally Malus) causes leaf to
curl downwards forming a bag-like gall, coloured yellowish-green tinged with
red. Alatae (BL 2.6-2.9 mm, with a yellowish-green abdomen) fly in late June-
early July. The secondary host is uncertain; roots of Larix according to Shinji
(1932; possibly due to confusion with another species), or of Rumex crispus
846 Prociphilus

according to Mordvilko (1935). Hori (1938) also noted that the secondary host
'may probably be Rumex'. Sexuparae occur in October-November, but have
not been described. In Japan, Korea and eastern Siberia.

Prociphilus laricis Shinji Only alate sexuparae described, BL c. 1.6mm,

found swarming out of the ground along the roots and trunk of Larix
leptolepis in Japan in late October-early November (Shinji, 1941). Primary
host unknown. [Sexuparae of a similar but undescribed species, with more
secondary rhinaria on ANT VI, have been collected from roots of Pinus
spp. (densiflora, thunbergii) in Japan (BMNH colln, leg. DHRL).]
Prociphilus ligustrifoliae (Tseng and Tao) Spring populations in curled
leaves of Ligustrum japonicus in China (Tseng and Tao, 1938). The globular
yellowish-white fundatrices, covered in white cottony wax, develop on the
buds in March-April and cause economic damage to trees used in rearing
the coccid Ericerus pela for commercial wax production in Sichuan (Zheng,
1981). The alatae produced in the leaf nests have BL c. 3.75mm, dark-
bordered wing veins, a pale green abdomen and secondary rhinaria distributed
III 32-36, IV 13-16, V 12-16, VI 3-7. They presumably fly to an unknown
secondary host. This species is closely related to, and possibly synonymous
with, P. osmanthae.
Prociphilus micheliae Hille Ris Lambers Only spring migrant alatae are
described; appearance in life unknown, BL 3.3-4.0 mm, secondary rhinaria
distributed III 28-35, IV 8-13, V 0, VI 0. In curled leaves of Michelia cham-
paca in northern India (Kashmir, West Bengal, Uttar Pradesh), becoming
mature in April-June (BMNH colln). Other morphs and life cycle unknown.
2n = 14 (Kar et al., 1990).

Prociphilus oleae Leach ex Risso Fundatrices and their progeny form com-
pact spring colonies covered in white wax wool on shoots near base of
trunk of Olea europea. Also recorded from Phillyrea media. Unusually for
a Prociphilus, no leaf deformation is caused. Progeny of fundatrix all alate,
BL c. 4 mm with smoky yellowish abdomen, migrating in May. The life
cycle is unknown, but apterae described from roots of Vitis vinifera in Sicily
(Barbagallo and Stroyan, 1982) could be the secondary host form of this
species. In Mediterranean area (France, Italy, Greece, Turkey). Roberti and
Monaco (1987) give a recent account.

Prociphilus oriens Mordvilko Fundatrices and their progeny infest young

growth of primary hosts in spring causing curling and clustering of leaves into
leaf-nest galls. Typical primary hosts are Fraxinus and Syringa, but there are
records of spring populations on a remarkable range of other plants, not only
in Oleaceae but including Rosaceae (Malus, Prunus, Sorbus) as well as
Lonicera, Salix, Tilia and Ulmus (Kono, 1940). Progeny of fundatrix are all
alatae, BL 3.5-5.5 mm, with yellowish-brown to dull reddish-green abdomen
covered with white wax; migrating in June-early July to roots of Abies
sachalinensis (incl. var. mayriana). Records from Picea roots need confirma-
 Prociphilus 847

tion. Return migration of sexuparae occurs in September-October (Hori,

1938). Recorded from India, China, eastern Siberia, Korea and Japan (type
locality). Yamaguchi (1985) studied sex ratio variation in this species.

Prociphilus osmanthae Essig and Kuwana Alatae developing in spring

colonies on Osmanthus spp. have BL 3.3-4.0 mm, wax-covered olive green
abdomen and dark-bordered forewing veins (Essig and Kuwana, 1918;
Moritsu, 1983). Secondary rhinaria of paratypes are distributed III 27-31, IV
9-12, V 8-12, VI 0. They migrate in late May to an unknown secondary host.
In Japan, and also reported to occur in Meghalaya, India (A.K. Ghosh and
Raychaudhuri, 1968c, on an unidentified host; Khuda-Bukhsh and Kar, 1990,
on Croton sp.). A.K. Ghosh (1984b) included various Japanese specimens
collected by Takahashi in his redescription of osmanthae; emigrant alatae
from Ligustrum, apterous exules from Michelia and alate sexuparae (with an
anocaudal process) from Osmanthus. It is not certain that all these forms are
the same species. Prociphilus ligustrifoliae also has dark-bordered wing veins
and may be a synonym, although it has more rhinaria, including some on ANT
VI. 2n = 18 (for aphids collected on Croton; Khuda-Bukhsh and Kar, 1990).

Prociphilus pergandei Smith Fundatrices curl leaflets of Fraxinus americana

in eastern USA. Progeny are all alate; BL 2.7-3.2 mm, covered with long
flocculent wax secretion, flying in June to an unknown secondary host (Smith,
1974a).

Prociphilus (Stagona) piceaerubensis Smith Apterae white to pale yellowish

with a tuft of wax on abdomen; BL 0.9-1.2 mm. On roots of Picea rubens,
inhabiting individual wax-lined cells associated with mycorrhizae, especially
in humus or where roots have penetrated decaying logs. Sexuparae occur in
September-October. Described from North Carolina, USA (Smith, 1969);
sexuparae which may be this species have also been trapped in Germany and
England (BMNH colln) and it is perhaps the form described as xylostei
'forma minima' by Tüllgren (1909).

Prociphilus (Stagona) pini Burmeister, Plate 6a, b Fundatrices and their

progeny curl and yellow the leaves of Crataegus spp. in spring. Progeny are
all alate; BL 2-3 mm, with light green to greyish-green abdomen, flying in
June-August to found wax-covered colonies on fine roots of Pinus sylvestris
and its close relatives; also recorded on P. contorta in UK (Danielsson and
Carter, 1993b). Sexuparae return to Crataegus in October-November, but
populations also persist throughout the year on Pinus roots. In northwest and
Central Europe. Smith (1969) gave distinguishing characters from the closely-
related North American species crataegistrobi. 2n = 16.

Prociphilus piniradicivorus Smith Apparently monoecious on Pinus spp.

(Smith, 1969). Apterae (presumably wax-covered in life, BL c. 2mm, with
protruding dark CAUDA and anal plate) live on roots of Pinus spp., attended
by ants and often in association with the very similar but host-alternating
848 Prociphilus

species P. carolinensis. Sexuparae (BL 2.9-3.3 mm) are produced in October-

November and fly only a short distance, depositing sexual morphs on trunks
of nearby pine trees. Colonies were experimentally reared from fundatrices
that hatched from eggs transferred to seedling P. taeda (Smith, 1969). In USA
(North Carolina, Maryland, Indiana).

Prociphilus (Pulvius) probosceus (Sanborn) Monoecious on Fraxinus.

Fundatrices and their developing alate progeny curl and distort leaves and
terminal growth of Fraxinus spp. in spring. More commonly found are the
very large apterae (BL 4.5-5.3 mm), in ant shelters at bases of water shoots
and trunks of young trees, at or below soil level (Smith and Stroyan, 1972).
Sexuparae are produced in October-November, but underground colonies
persist throughout the year. In eastern North America.

Prociphilus sasakii Monzen Described from Malus pumila in Japan

(Monzen, 1927, 1929). Large, globular fundatrices (BL c. 3.3mm) and their
developing alate progeny, covered with white woolly wax, cause apple leaves
to become curled, bloated and yellowed in spring. Alatae, BL c. 3.1 mm with
pale green abdomen and secondary rhinaria distributed ANT III 15-18, IV
5-8, V 5-7, VI 0, migrate to an unknown secondary host. (See also discussion
of this species under P. crataegicola.)

Prociphilus taxus Ghosh, Chakrabarti, Chowdhuri and Raychaudhuri Only

known from the alate sexupara, collected on roots of Taxus baccata in
Himachal Pradesh, India (A.K. Ghosh et al., 1969b); BL 2.9-3.5 mm, secon-
dary rhinaria distributed ANT III 18-22, IV 8-10, V 6.10, VI 2-5). The anal
plate projects as a collar around the CAUDA; a similar anocaudal process is
known in sexuparae of certain other Prociphilus (keyed by A.K. Ghosh,
1984b, p. 287). Life cycle unknown.

Prociphilus ushikoroshi Shinji Fundatrices and their developing alate pro-

geny feed on leaves of Photinia (= Pourthiaca) villosa in spring, causing edges
of leaves to curl towards underside and become thickened and yellowish-green
(Monzen, 1929; as P. pourthiacae). Alatae (BL c. 2.1mm, with yellowish-
green abdomen covered in white woolly wax and secondary rhinaria distri-
buted ANT III 10-14, IV 4-7, V 1-4, VI 0-1) migrate in June. The secondary
host is uncertain, possibly roots of Larix (Shinji, 1932). Sexuparae return to
deposit sexual morphs on trunk of Photinia in October (Monzen, 1929). In
Japan and Korea. [In India, alatae with many more secondary rhinaria (III
35-36, IV 9-10, V 10-11, VI 0-2) have been collected on Photinia notaniana
(A.K. Ghosh et al., 1970; Prociphilus 'sp. A').]

Prociphilus (Stagona) xylostei (De Geer) Fundatrix and developing alate

progeny live in copious wax on Lonicera spp. in spring, causing leaves to
curl and become spotted with yellow (Mordvilko, 1935; Heie, 1980). Alatae
(BL 2.3-3.7 mm, with pale green abdomen) complete their development on the
twigs in late June-July and migrate to found wax-covered colonies on roots
 Protopterocallis 849

of Picea abies. Sexuparae return to Lonicera in October-November. Widely

distributed in Europe, in India (Uttar Pradesh, Kashmir) and introduced to
North America. Differences from P. piceaerubensis were discussed by Smith
(1969). Populations in Siberia are regarded as a subspecies, P. xylostei
ussuricus Pashchenko, 1988b (= P. xylostei ssp. orientalis Mordvilko, 1935;
nec orientalis Mordvilko, 1929a). 2n = 10 (Pal and Khuda-Bukhsh, 1983).

PROTOHORMAPHIS Shaposhnikov and Gabrid

Hormaphidinae(?): Hormaphidini(?)

One conifer-feeding species of uncertain phylogenetic position, possibly

related to Tsugaphis. Only the apterous vivipara is known.

Protohormaphis piceae Shaposhnikov and Gabrid Apterae oval, slightly

flattened, dark green; BL c. 1.0mm. Immatures are lighter bluish-green with
white wax markings. Found feeding on needles of Picea pungens in May-June,
in Kirgizia (Shaposhnikov and Gabrid, 1987). Life cycle unknown.

PROTOPTEROCALLIS Richards Drepanosiphinae: Phyllaphidini

Four eastern North American species on Carya related to Monellia and

Monelliopsis, but larger and more robust, and with a shorter processus
terminalis. All adult viviparae are alate. Bissell (1978) reviewed the genus.
All stages are said by Bissell to feed on upper surface of leaf, usually along
mid-ribs of basal leaflets.

Protopterocallis fumipennella (Fitch) (= P. canadensis Richards) Alatae

broad-bodied, abdomen yellow-orange, yellowish-green or olive green dusted
with wax, prothorax darker, dorsal abdominal tubercles brown-tipped; BL
2.0-2.2 mm. Singly or in small colonies on undersides(?) of leaflets of Carya
spp., especially cordiformis (but not illinoensis), in eastern USA and Canada
(Quednau, 1966; Bissell, 1986). Monoecious holocyclic; oviparae and males
in September-October (Bissell, 1978).

Protopterocallis gigantea Bissell Alatae broad-bodied, abdomen dull yellow

with irregular streaks of orange or red, dorsal abdominal tubercles brown-
tipped and forewing veins (especially Cu lb ) dark brown; BL 2.5-3.0 mm.
Monoecious holocyclic on Carya spp. (but not illinoensis) in eastern USA
(Washington, DC, Maryland, Georgia). Oviparae and males in October
(Bissell, 1978). 2n = 10.
850 Protrama

Protopterocallis pergandei Bissell Appearance in life unknown; alata broad-

bodied, BL 2.5-2.9 mm. Type material from Carya sp. in Florida, USA.
Monoecious holocyclic; an ovipara was collected on Carya pallida in Georgia
in November (Bissell, 1978).

Protopterocallis quadrata Bissell Alata broad-bodied, yellowish; BL 1.7-

2.8mm (Bissell, 1978). On Carya tomentosa and Carya sp. in eastern
USA (Pennsylvania, Washington, DC, Georgia). Biology and sexuales not
recorded.

PROTRAMA Baker Lachninae: Tramini

About ten species mostly on roots of Compositae (see B & E, 1984, p. 332).

Protrama tamaricis (Nevsky) Described from a single alata on roots of

Tamarix ramosissima in Uzbekistan (Nevsky, 1951, as Eulachnus tamaricis),
but P. xerophilaphica, apterae and alatae of which were described from the
same host in Kazakhstan (Juchnevitch and Kan, 1971), is probably a synonym.
Creamy-white aphids, BL 3.9-4.7 mm, attended by ants. Mamontova (1991)
provided a redescription and discussed its synonymy and generic affiliation.
Czylok (1990) placed P. xerophilaphica in Eotrama.

PSEUDASIPHONAPHIS Robinson Aphidinae: Aphidini

One North American species with very small flangeless siphunculi.

Pseudasiphonaphis corni (Tissot) Spring colonies, on undersides of leaves

and along young shoots of Cornus microcarpa in Florida, contained brown
apterae (BL 1.7-2.1 mm) underneath an extensive light greyish-white woolly
wax secretion. Alatae have dark wing veins and a reddish-brown abdomen
(Tissot, 1929). Further north in the USA (Pennsylvania, Illinois, New York),
there are very few records from Cornus and colonies of this species occur most
commonly on Pilea pumila (Urticaceae). A colony has also been found on a
labiate, Scutellaria laterifolia (Robinson, 1965). Host alternation is indicated,
but requires confirmation; there are no records of the return migration to
Cornus, or of sexual morphs. In Florida, anholocyclic populations persist all
year on Cornus (Tissot, 1929).
 Pseudoregma 851

PSEUDESSIGELLA Hille Ris Lambers Lachninae: Cinarini

One pine-feeding species in Pakistan related to Eulachnus and Essigella;

Sorensen (1990) discussed its relationship to these other genera.

Pseudessigella brachychaeta Hille Ris Lambers Apterae pale green, narrowly

elongate; BL 2.0-2.6mm. On needles of Pinus griffithii (= wallichiana) in
Pakistan (Hille Ris Lambers, 1966c). Other morphs and life cycle unknown.

PSEUDOPTEROCOMMA MacGillivray Chaitophorinae

Two poplar-feeding species only known from Canada and northwest USA,
related to Chaitophorus but with a distinctive, long, thick processus terminalis
clothed with hairs like the rest of the antenna, and pore-like siphunculi.
Richards (1972a) provided a key.

Pseudopterocomma canadensis Richards Apterae grey or black; BL 3.0-

3.5mm. Alatae have dark-bordered wing veins. On Populus tremuloides in
Canada (Manitoba, Ontario, Saskatchewan). The feeding position on the
plant is not recorded, but is likely to be low down on the stem or suckers,
or possibly below ground level. Sexual morphs and life cycle unknown.

Pseudopterocomma hughi (MacGillivray) Apterae cream-coloured; BL 2.3-

3.0mm. Alatae have dark-bordered wing veins. On roots of Populus tremu-
loides, attended by ants. In Maine, USA (BMNH colln, leg. G. Remaudière)
and Canada (Manitoba, New Brunswick, Quebec). Monoecious holocyclic,
with fundatrix in May and apterous males and oviparae in September
(MacGillivray, 1963).

PSEUDOREGMA Doncaster Hormaphidinae: Cerataphidini

About 12 species of wax-dusted aphids living mostly on either bamboos or

Zingiberidaceae, with a few species on other Gramineae and some other
monocotyledons in Southeast Asia. Most of the species were originally
described in Oregma and may be found in accounts of Oregma until 1966. The
long-suspected host alternation to Styrax was recently confirmed for two
852 Pseudoregma

species in Taiwan (Aoki and Kurosu, 199la). There is a remarkable poly-

morphism in the first or second instar, some nymphs being pseudoscorpion-
like and functioning as soldiers for the colony (Aoki, 1987). Accounts of
the bamboo-feeding species are available for Taiwan (Liao, 1976), Java
(Noordam, 1991) and India (A.K. Ghosh, 1975a).
Pseudoregma alexanderi (Takahashi) Apterae rather elongate pear-shaped,
pale brown to brownish-black with a purple tinge, lightly dusted with wax
powder that is often thicker on the posterior abdomen; BL 2.2-4.3 mm. Large
colonies attack the young shoots and leaves of bamboos (Dendrocalamus,
Miscanthus) in Taiwan, Nepal and northeast India. Soldier behaviour was
studied by Aoki et al. (1981). Life cycle is uncertain; alate sexuparae are
produced in May in Taiwan, but the primary host is unidentified and popula-
tions also persist throughout the year on Dendrocalamus (Aoki et al., 1981).
M.R. Ghosh, Pal and Raychaudhuri (1977b) erected a new genus,
Paraoregma, for this species. 2n = 12 (Khuda-Bukhsh and Kar, 1987).

Pseudoregma bambusicola (Takahashi) (= albostriata Liao?) Galls on

Styrax suberifolia in Taiwan consist of clusters of elongate pod-shaped sub-
galls with petiolate bases (cf. P. koshuensis) and apical openings, arising from
a single point on the stem (Aoki and Kurosu, 1992). Gall populations include
second instar soldier larvae. Alatae emerge from galls in July-September
to colonize bamboos and alate sexuparae return in December. Apterae on
bamboos are broadly oval, in life grey-brown or dark purplish to greenish-
brown or black, dusted with granular wax; BL 1.5-3.6 mm. Individuals in
young colonies may have 3 longitudinal rows (2 marginal, 1 spinal) of white
wax tufts, but in older colonies these are less developed and adults have a
thick cushion of wax posterior to the siphunculi (Noordam, 1991). Dense,
ant-attended colonies occur on the shoots, young stems and leaf bases of
Bambusa spp., especially B. multiplex. Also recorded from Dendrocala-
mus giganteus. Found in Japan, Ryukyu, Taiwan, Hong Kong, Thailand,
Indonesia, Malaysia, India, Sri Lanka and Fiji. Sunose et al. (1991) studied
population dynamics of P. bambusicola on B. multiplex in Japan, Arakaki
and Yoshiyasu (1988) studied predation by a pyralid moth and Ohara (1985)
studied defensive behaviour of soldiers against syrphid larvae. In Japan,
parthenogenetic reproduction continues throughout the year; sexuparae are
produced in autumn (Sunose et al., 1991), although the primary host is not
available. Noordam (1991) regarded P. albostriata Liao, 1976, described from
B. oldhami and D. giganteus in Taiwan, as a synonym, although in life this
species was clearly described as having 4 longitudinal white stripes, not 3 as
in P. bambusicola. Pseudoregma cantonensis Takahashi, differentiated by
having greater sclerotization of the dorsum, is another possible synonym.
2n = 12 (Chen and Zhang, 1985b).
Pseudoregma dendrocalami (Takahashi) Green to brownish-green apterae
with four wax stripes, described from Dendrocalamus flagellifer in Sumatra
(Takahashi, 1935a). Possibly P. albostriata is this species.
 Pseudothoracaphis 853

Pseudoregma koshuensis (Takahashi) (= Astegopteryx shitosanensis Taka-

hashi) Galls on Styrax suberifolia in Taiwan are grey-brown, hard, banana-
or flask-shaped, about 28 mm long with apical opening, occurring in bunches
(Fig. 124L, p. 578). Their inhabitants include many pseudoscorpion-like
second instar soldiers (Aoki, 1982). Heteroecious holocyclic; alatae migrate in
June to Bambusa spp. (Aoki and Kurosu, 1991b). Apterae infesting the culms
of bamboo plants are broadly oval, purplish to dark brown, covered with
white wax; BL 1.6-2.0 mm (Takahashi, 1924a). Individuals on leaves are
generally darker, more dorsoventrally flattened, and the wax forms marginal
tufts on posterior abdominal segments; they were described as a distinct
species, pseudomontana (Takahashi, 1924a), but most authors (e.g. Aoki and
Kurosu, 1991b) now regard the two forms as conspecific. Parthenogenetic
populations may also persist through the year on bamboos.

Pseudoregma montana (van der Goot) Apterae broadly oval, yellowish to

violet-coloured with yellowish head and prothorax, dusted with wax and with
4 variably developed longitudinal rows of small white wax tufts (2 spinal, 2
marginal); BL 1.9-2.8 mm (Noordam, 1991). Immatures are bright yellow to
orange, dusted with wax. In dense colonies on undersides of leaves near the
leaf bases of unidentified bamboos in Java. Large populations spread over the
whole shoots. Apart from its size and colour, the difference between this
species and P. koshunensis from Taiwan is unclear.

Pseudoregma pendleburyi (Takahashi) Apterae blackish-brown, marbled;

BL 1.3-2.7 mm. Originally described from Malaya on Bambusa sp.
(Takahashi, 1950) and redescribed (including first instar soldiers) from Java
by Noordam (1991). Also recorded from West Irian (BMNH colln, leg. E.W.
van Heurn). The populations in Java were living on the culms, but bore a close
resemblance in size, colour and development of wax glands to the leaf-feeding
population in Malaya that Takahashi (1950) described as a distinct species,
gombakana. Development of wax glands in Pseudoregma depends greatly
on age of colony and degree of alatiformity, and it seems probable that
gombakana is a junior synonym of pendleburyi, which was described first in
Takahashi's (1950) paper. Oregma uscare Tao, described from unidentified
bamboos in Okinawa (Tao, 1964), is possibly also a synonym. Life cycle is
unknown.

PSEUDOTHORACAPHIS Raychaudhuri, Ghosh and Das

Hormaphidinae: Nipponaphidini

One species on Quercus in northern India, perhaps related to Allothoracaphis

but lacking marginal prosomal hairs.
854 Pterasthenia

Pseudothoracaphis himachali Raychaudhuri, Ghosh and Das Apterae

oval, dark brown, dorsoventrally flattened, strongly sclerotized; BL 1.1-
1.6mm. On undersides of leaves of Quercus incana and Quercus sp. in India
(Himachal Pradesh, Uttar Pradesh), attended by ants. The host plant of the
holotype was reported to be Ilex sp., but this is likely to be an error. All
specimens were collected in October (Raychaudhuri et al., 1980a); other
morphs and life cycle unknown. Heminipponaphis querciphaga Chakrabarti
and Raha (1985) seems to be a synonym.

PTERASTHENIA Stroyan Drepanosiphinae: Phyllaphidini

Four African species related to Paoliella. Remaudière and Autrique (1985)

discussed the taxonomie status of the genus. One species occurs on Milletia;
the others are on herbaceous Leguminosae.

Pterasthenia matileae Remaudière and Quednau All viviparae alate, pale

yellow with bluish-white cottony wax on body, legs and antennae; BL 1.5-
2.3 mm. Living all year on Milletia eetveldiana in the Congo Republic;
colonizing undersides of leaves in dry season, but in wet season found
especially on young shoots at base of stem (Remaudière and Quednau, 1988).
Parthenogenetic reproduction continues throughout the year and no sexual
morphs are known. This aphid is attacked by a specific parasitoid (Trioxys
pterastheniae; Starý et al., 1977).

PTEROCALLIS Passerini Drepanosiphinae: Phyllaphidini

Fourteen species of small pale aphids with a short antennal terminal process
living on Betulaceae, especially Alnus. They are usually dispersed on under-
sides of leaves and are mostly not attended by ants. Apterous viviparae occur
more commonly than in most Drepanosiphinae, particularly in Pterocallis
s. str. which has 4 species in North America, 3 in Europe and 3 in East Asia.
The subgenera Paratinocallis (1 sp.) and Recticallis (3 spp.) are wholly East
Asian. Mesocallis is here considered as a separate genus. Accounts are available
for Britain (Stroyan, 1977), Fennoscandia and Denmark (Heie, 1982), Central
Europe (Börner, 1949), Japan (Higuchi, 1972), India (Chakrabarti, 1988;
A.K. Ghosh and Quednau, 1990), USA (Granovsky, 1928a; as Myzocallis)
and Canada (Richards, 1965).

Pterocallis affinis Chakrabarti Alatae pale yellow, with antennae and legs
mainly pale; BL 1.5-2.0 mm. On Corylus colurna in Uttar Pradesh, India. No
 Pterocallis 855

apterae are recorded. Monoecious holocyclic; fundatrices in April (Banerjee

and Chakrabarti, 1991), oviparae and males in October (Chakrabarti, 1988).

Pterocallis albida Börner Apterae and alatae are yellowish-white or greenish-

white, shining, with pale antennae and legs; BL 1.0-1.2 mm. On Alnus incana,
dispersed on undersides of leaves. Widely distributed in continental Europe
(see Lampel, 1983 for details). Monoecious holocyclic; oviparae and males in
September (Italy; BMNH colln, leg. H.L.G. Stroyan).

Pterocallis alni (De Geer) Apterae and alatae are yellowish-white to yellowish-
green, with antennae ringed with black, a black spot near apex of hind femur
and black tarsi; BL 1.3-2.0 mm. Almost exclusively on Alnus glutinosa in
Europe, living dispersed on undersides of leaves and only rarely attended by
ants. Common and widespread in Europe, east to Turkey, Iran and Lebanon;
introduced to North America (Essig, 1912b), New Zealand (Lowe, 1968),
Australia (Carver, 1980a) and Chile (Carrillo, 1980). In North America it
seems able to feed on various native Alnus spp. (BMNH colln). South-
west Asian populations colonize A. orientalis and A. subcordata as well as
A. glutinosa and tend to be intermediate in morphology between P. alni and
P. maculata, so their specific identity is often uncertain. Monoecious holo-
cyclic, with oviparae and alate males in September. Gange and Pryse (1990)
investigated the effects of temperature and nutrition on development, and
Gange and Llewellyn (1988) studied egg distribution and mortality. Pterocallis
alni on alder windbreaks may be a useful alternative food for predators of
orchard pests (Gange and Llewellyn, 1989). 2n = 20.

Pterocallis alnifoliae (Fitch) Apterae pale green, pale yellow or whitish with
darker green irregular patches (Granovsky, 1928a); BL 1.1-1.5 mm. Alatae
have pale greenish-yellow abdomen and darker yellow to orange head and
thorax; BL 1.3-1.7 mm. On undersides of leaves of Alnus rugosa and A.
serrulata in eastern North America. Monoecious holocyclic; oviparae and
alate males in September. [Granovsky (1928a) described a very similar but
smaller species, A. rhombifoliae, from Alnus rhombifolia in California and
also from A. incana (= rugosa) in Wisconsin. Collections from Maine on
Alnus sp. (BMNH colln, leg. H.L.G. Stroyan) and Quebec on A. rugosa
(BMNH colln, leg. DHRL) seem to be this species. However, it is not clear
how apterae of rhombifoliae can be distinguished from those of alnifoliae.
According to F.W. Quednau (pers. comm.), P. alnifoliae has only alate
viviparae and all apterae are rhombifoliae. Further work is needed to clarify
the situation; in our key to Alnus-feeders, both species are therefore taken out
in two places, but discriminated only on the basis of characters of alatae.]

Pterocallis (Recticallis) alnijaponicae (Matsumura) All adult viviparae alate,

yellowish with pale antennae and legs, abdomen with marginal rows of brown
spots and spinal tubercles sometimes also brown-tipped; BL 1.5-2.1mm
(Higuchi, 1972). Recorded from various Alnus spp. in Japan and Korea.
Sexual morphs undescribed, life cycle unknown.
856 Pterocallis

Pterocallis (Paratinocallis) corylicola (Higuchi) Adult viviparae all alate,

pale yellow with usually only ANT V and VI, apices of tibiae and tarsi pale
brown; BL c. 1.5 mm. Described from Corylus sieboldiana in Japan (Higuchi,
1972) and also recorded from Korea (on C. heterophylla; Quednau, 1979).
Specimens from C. heterophylla from Yunnan Province, China have a longer
R IV+V and were described (Zhang and Zhong, 1985c) as ssp. yunnanensis
Zhang. Sexual morphs and life cycle unknown. 2n = 10 (Blackman, 1986).

Pterocallis heterophyllus Quednau Alatae whitish to very pale yellow with

blackish apices to antennal segments, a black spot at base of pterostigma,
narrowly brown-bordered forewing veins and brown tips to dorsal body hairs;
BL c. 1.6-1.7 mm. Apterae not known. Described from specimens collected
on Corylus heterophylla in Korea in August and November (Quednau, 1979).
Sexual morphs and life cycle unknown.

Pterocallis maculata (von Heyden) Apterae yellowish-green or green with a

variably developed pattern of rather diffuse dark green dorsal cross-bands,
black apices to antennal segments, a black spot near apex of hind femur and
black tarsi; BL 1.4-2.1 mm. Alatae have greenish abdomen with dark green
markings. On undersides of leaves of Alnus glutinosa, usually along main
veins, usually attended by ants (cf. alni). Widely distributed in Europe, east
to Turkey and Iran. Monoecious holocyclic; apterous males and oviparae in
early September (Stroyan, 1955).

Pterocallis montana (Higuchi) Alatae pale yellow, with pale antennae,

wings and legs; BL 0.8-1.2 mm. Apterous viviparae unknown and probably
do not occur. On undersides of leaves of Corylus sieboldiana (Higuchi, 1972;
as Myzocallis montana) and Carpinus japonica (BMNH colln, leg. RLB) in
Japan. Sexual morphs and life cycle unknown. 2n = 16 (Blackman, 1986).

Pterocallis (Recticattis) nigrostriata (Shinji) All adult viviparae alate, yellow-

ish with black apices to antennal segments; a dark patch in middle of hind
femur; forewings characteristically pigmented with broad bands of fuscous
bordering veins and lines of darker pigment between veins near wing-tip; black
spinal tubercles on ABD TERG 1-4, and black marginal spots on ABD TERG
3-4; BL 1.4-2.0 mm. Immatures have conspicuous long dark dorsal hairs. On
undersides of leaves of various Alnus spp. in Japan (Takahashi, 1965b) and
Korea. Paik (1972) recorded oviparae in Korea in early November. 2n = c. 26*.

Pterocallis ostryae (Börner) Apterae and alatae yellow; BL 0.9-1.4 mm

(Börner, 1949). Alatae have a dark spot at apex of hind femur and a dark
triangle at base of Cu1b in forewing. On undersides of leaves of Ostrya
carpinifolia in alpine and southern Europe (Austria, France, Italy). Sexual
morphs and life cycle unknown.

Pterocallis (Recticattis) pseudoalni (Takahashi) All adult viviparae alate,

with pale brown head and thorax and yellow abdomen; antennae banded with
 Pterocomma 857

brown-black, forewing veins narrowly bordered with fuscous, dark hind legs,
dark finger-like spinal tubercle on ABD TERG 1, dark marginal spots on ABD
TERG 3-4 and black distal halves to siphunculi; BL 1.6-2.1 mm. On under-
sides of leaves of Alnus formosana in Taiwan, where it reproduces partheno-
genetically throughout the year (Takahashi, 1923; as Myzocallis pseudoalni).
Takahashi (1924a) provided detailed descriptions of all immature stages.
Granovsky (1928a) provided an illustrated redescription.

PTEROCHLOROIDES Mordvilko Lachninae: Lachnini

One species with a double row of large spinal tubercles.

Pterochloroides persicae (Cholodkovsky) Apterae shiny, oval, dark brown

to black; BL 2.7-4.2 mm. Living along undersides of larger branches and
sometimes forming large colonies on trunks of woody Rosaceae (especially
Prunus spp.). Large amounts of honeydew are produced and colonies are
regularly attended by ants. In southern Europe, North Africa, southwest and
Central Asia, India and Pakistan; it appears to be gradually extending its
range, with recent first reports of economically damaging populations in
Romania and Tunisia. Monoecious holocyclic on Prunus (persica, armeniaca,
spinosa) in cooler regions according to Archangelsky (1917); but males have
never been described and most populations seem anholocyclic. Ecological
studies have been carried out in Egypt (Darwish et al., 1989), Italy (Ciampolini
and Martorelli, 1978), Lebanon (Talhouk, 1977) and former Yugoslavia
(Velimirovic, 1976). 2n = 20.

PTEROCOMMA Buckton Aphidinae: Pterocommatini

About 30 species of robust, hairy aphids living on Salicaceae in the northern

hemisphere. They are evenly distributed between western palaearctic (9 spp.),
eastern palaearctic (12 spp.) and nearctic (8 spp.). They tend to be dark-
coloured, but with often contrastingly brightly coloured siphunculi and their
colonies on the bark of branches and twigs are almost always attended by ants.
Szelegiewicz (1965) reviewed the genus and regional accounts are available for
Britain (Stroyan, 1984), Fennoscandia and Denmark (Heie, 1986), Central
Europe (Heinze, 1961b), eastern Russia (Pashchenko, 1988b), China (Zhang
and Zhong, 1980b), US Rocky Mountain region (Palmer, 1952) and Canada
plus Greenland (Richards, 1967). Specialized parasitoids include Aphidius
pterocommae and perhaps also Aphelinus fulvus, which is also recorded from
willow-feeding Chaitophorus.
858 Pterocomma

Pterocomma bailangense Zhang Colour in life unrecorded; BL of aptera

c. 4.6mm. On an unidentified Populus sp. in Tibet (Zhang and Zhong,
1981b). Other morphs and biology unknown.

Pterocomma beulahense (Cockerell) Apterae yellowish-olive green (Palmer,

1952), probably sometimes darker; BL 2.6-4.0 mm. Alatae have grey-brown
abdomen with black dorsal cross-bands. On twigs of Populus tremuloides
in the Rocky Mountains, from Washington to New Mexico. Presumably
monoecious holocyclic; a male was collected in September (BMNH colln, leg.
DHRL). Richards (1967) synonymized beulahense, which seems to be specific
to P. tremuloides, with the more polyphagous P. bicolor; the two can be
distinguished by the characters given in the poplar-aphid key.

Pterocomma bicolor (Oestlund) Apterae reddish-brown to dusky greenish-

yellow, greyish or bluish-green, with yellow-orange siphunculi; BL 3.2-4.8 mm.
Alatae have reddish-brown to dusky-green abdomen with variably-developed
dark brown to black dorsal markings, only sometimes forming complete cross-
bands. On bark of twigs and small branches, or on trunks of saplings, where
they occasionally feed just below ground level. Native host plants are Populus
spp. (especially angustifolia, balsamifera, tremuloides, trichocarpa), but this
species also occurs commonly throughout North America on naturalized
Salix spp. (alba, babylonica). Widespread in North America and in Mexico.
Monoecious holocyclic; oviparae and alate males in October-November.
2n = 8.

Pterocomma (Stauroceras) chaetosiphon (Börner) Apterae greenish-grey to

brownish, with yellow siphunculi; BL 2.5-3.0 mm. Alatae not described. In
bark crevices of trunk and branches of Salix spp. (fragilis, alba), only known
from central Germany (Heinze, 1961b). Monoecious holocyclic; oviparae in
late September (BMNH colln, leg. DHRL).

Pterocomma groenlandica Hille Ris Lambers Colour in life unknown; BL

of aptera 2.8-3.7 mm. Alatae have well-developed dark dorsal abdominal
cross-bands. On Salix sp. in Greenland. Monoecious holocyclic; oviparae and
alate males in late July-August (Hille Ris Lambers, 1952). Closely related
to P. bicolor, of which it was placed as a subspecies by Richards (1967).
Pterocomma baicalense, described from osiers in Siberia (Ivanovskaya, 1976),
is very similar and could possibly be a synonym.

Pterocomma italica Barbagallo and Stroyan Apterae very dark brown, with
a fine spinal line of whitish wax crossed by two transverse wax bands on about
ABD TERG 1 and 5 and orange-yellow siphunculi; BL 2.3-3.4 mm. Alatae
with variable dorsal abdominal markings, not forming cross-bands. On twigs
or at base of stems of Salix purpurea, in mountainous areas of Italy and
Switzerland (Barbagallo and Stroyan, 1978). Sexual morphs and life cycle
unknown. Very similar to P. jacksoni. Heavy parasitization by an Aphidius
species is reported in the original description.
 Pterocomma 859

Pterocomma jacksoni Theobald Apterae varying in colour from pinkish or

reddish-brown to brown-black, with a greenish tinge, sometimes with fine lines
of wax as in italica and salicis and orange-yellow siphunculi; BL 2.2-4.3 mm.
Alatae without dorsal abdominal cross-bands. On branches or stems of Salix
spp. (aurita, caprea, cinerea, repens), forming dense ant-attended colonies
either above or below ground level. Widely distributed in Europe (see Heie,
1986), but uncommon. Sexual morphs and life cycle unknown. 2n = c. 30-34
(Kuznetsova and Shaposhnikov, 1973).

Pterocomma kozchukovae Pashchenko Apterae orange-brown, brown or

brownish-green, dusted with wax; BL c. 3.5 mm. Forming dense ant-attended
colonies on stems and suckers of Populus bolleana in maritime eastern Russia.
Monoecious holocyclic, with alate males (Pashchenko, 1988b).

Pterocomma lhasapopuleum Zhang Colour in life not recorded; BL of

aptera c. 3.7mm. Alatae have dark dorsal abdominal cross-bands. On an
unidentified Populus sp. in Tibet (Zhang and Zhong, 1981b). Sexual morphs
and life cycle unknown.

Pterocomma pilosum Buckton Apterae greyish or brownish, with yellow

siphunculi; BL 2.3-4.0 mm. Alatae have dark dorsal abdominal cross-bands,
sometimes divided medially into paired patches. On twigs and branches or in
bark crevices of numerous Salix spp., usually attended by ants. In northwest,
Central and Eastern Europe and southwest Asia (Iran); introduced to western
North America (Utah, British Columbia). Monoecious holocyclic, with
oviparae and alate males in October. A closely-related form commonly
regarded as a subspecies, P. pilosum konoi Hori, is recorded from an even
wider range of Salix spp. and occurs in Europe and eastward across Asia to
Japan and Korea. Oviparae of konoi have darker, more swollen hind tibiae
than those of pilosum s. str., bearing many more scent plaques (Heie, 1986,
citing Danielsson). In Europe, konoi may occur in mixed colonies with
pilosum s. str., indicating that there may be an effective isolating mechanism
and that the two taxa are probably distinct species. Szelegiewicz (1967)
described another variant population, from Salix alba var. pendula in Poland,
as a subspecies P. pilosum sarmaticum; similar specimens were also found
on S. alba in The Netherlands (BMNH colln, leg. H.L.G. Stroyan). [Moritsu's
(1983) account and illustration of an aphid with long fine hairs on the
siphunculi under the name pilosum must refer to some other, apparently
undescribed, Japanese species.] 2n = 8.

Pterocomma populeum (Kaltenbach), Plate l0e, f Apterae yellowish-grey or

brownish, with paired dark patches at least on posterior abdominal tergites,
intersegmental cross-bands of greyish-white wax and pale siphunculi; BL
2.7-4.3 mm. Alatae have broad dark dorsal abdominal cross-bands. On
branches or two-year-old twigs, or in bark crevices, of many Populus spp.,
but especially common on P. nigra and its varieties. Widespread in Europe,
including Spain, North Africa, southwest Asia, and east to Siberia and
860 Pterocomma

Mongolia. Introduced to North America (see Smith and Parron, 1978; Forbes
and Chan, 1986a) and South America (Argentina; Chiesa Molinari, 1942, and
Chile; BMNH colln, leg. E. Zuniga). Monoecious holocyclic; oviparae and
alate males occur in October-November. Martin-Bernal and Padro Simarro
(1986) reviewed natural enemies. Börner (1950) described a form, dubium,
from P. alba in Germany, resembling populeum but differing in colour (silver
grey) and absence of marginal tubercles; Holman and Szelegiewicz (1974a)
reported similar aphids (as P. populeum dubium) from Mongolia, collected
on P. laurifolia. [There are numerous records of this species from Salix, but
these are almost certainly due to confusion with other species (pilosum and
pilosum ssp. konoi in palaearctic; ?sanguiceps in North America.]

Pterocomma populifoliae (Fitch) Apterae pinkish to brown, variably dusted

with greyish wax, with siphunculi pale to dusky orange (Richards, 1967);
BL 2.2-3.8 mm. Alatae with variably developed dorsal dark patches, not
forming cross-bars except on posterior tergites. On Populus spp. (balsamifera,
grandidentata, tremuloides) in northeastern USA and Canada, usually col-
onizing roots or at bases of stems of sapling trees. Generally confused in the
North American literature with the widely distributed and more polyphagous
P. bicolor (Richards, 1967); Palmer's (1952) account of populifoliae should
be referred to that species. The identity of populations frequently recorded
as P. populifoliae on Populus ciliata in India (e.g. Saha and Chakrabarti,
1988d) needs confirmation, as it seems more likely that there is a similar,
undescribed Indian species on this host. Monoecious holocyclic in North
America, with oviparae and alate males in September.

Pterocomma pseudopopuleum Palmer Apterae greyish to yellowish-olive-

brown, finely dusted with wax; BL 3-4 mm. Alatae have broad dark dorsal
abdominal cross-bands. On bark of twigs and small branches of Populus
spp. (especially angustifolia and deltoides var. occidentalis), in the Rocky
Mountain region (Idaho, Utah, Colorado, New Mexico). Monoecious holo-
cyclic; oviparae and alate males in October-November (Palmer, 1952).

Pterocomma rufipes (Hartig) (= steinheili Mordvilko) Apterae variably

pigmented, grey or dull reddish-brown to dark brown with spots of powdery
wax and yellowish siphunculi; BL 2.9-4.6 mm. Alatae have variably developed
dorsal abdominal cross-bands. On twigs and young branches of numerous
Salix spp., attended by ants. Heie (1986) recorded it also on ornamental and
hybrid balsam poplars. In northwest and Central Europe, Iceland, Mongolia,
Siberia (Pashchenko, 1988b), and introduced to Canada (Manitoba; BMNH
colln, leg. A.G. Robinson). Oviparae and alate males in September. Larger,
darker specimens with longer siphunculi are sometimes regarded as a separate
species, P. ringdahli, but are most probably part of a range of intraspecific
variation (Heie, 1986). Pterocomma henanse Zhang, on Salix sp. in China
(Zhang and Zhong, 1980b) and P. tibetasalicis Zhang, on Salix sp. in Tibet
(Zhang and Zhong, 1981b), are closely related, if not synonyms. 2n = 8 or
9 (Kuznetsova, 1974, as steinheili).
 Pterocomma 861

Pterocomma salicis (Linnaeus) Apterae and alatae greenish-black to black,

marked dorsally with greyish-white wax powder including a spinal stripe and
lateral spots and bright orange siphunculi; BL 2.7-4.5 mm. Forming dense
colonies on two-year-old twigs and wands of Salix spp., attended by ants.
Widely distributed in Europe; Central Asia, Mongolia, eastern Siberia;
introduced and widespread in North America. Monoecious holocyclic, with
apterous males and oviparae in October-November. Holman and Szelegiewicz
(1974a) discriminated a subspecies, P. salicis rohdendorfi, on Salix sp. in
Transbaikalia and Mongolia. 2n = 58*.

Pterocomma salijaponica (Shinji) Apterae orange or brown with orange

siphunculi (Shinji, 1924, 1927); BL 2.8-3.5 mm. Alatae have dark dorsal
abdominal cross-bands, sometimes broken in midline. On stems of Salix
spp. (babylonica, gracilistyla, koriyanagi) in Japan. Monoecious holocyclic,
with alate males (Shinji, 1941). The identity of this species is uncertain; it
could not be included in the key to Salix-feeders on the basis of the available
information. It was listed in error as a Plocamaphis by Eastop and Hille Ris
Lambers (1976) and Paik's (1965) account under Clavigerus salijaponica is of
Plocamaphis coreana. The karyotype is also uncertain; 2n = 8 or 2n = 22
(Shinji, 1927, 1931, 1941 - see Blackman, 1986, as Plocamaphis salijaponica).

Pterocomma sanguiceps Richards Apterae have head and prothorax bright

red, rest of dorsum bluish, with yellow to pink siphunculi; BL 3.0-3.8 mm.
Alatae have abdomen brownish or bluish, without dark dorsal cross-bands.
On Salix spp. (babylonica, exigua) in British Columbia, Canada (Richards,
1967; BMNH colln, leg. C.K. Chan). 2n = 8*.

Pterocomma sanpunum Zhang Apterae yellowish-red; BL c. 3.9mm. On

Populus cathayana and P. simonii in China (Zhang and Zhong, 1980b). Other
morphs and life cycle unknown. Closely related to, if not synonymous with,
P. yezoense.

Pterocomma smithiae (Monell) Apterae dark reddish-brown with bluish

pruinosity and yellow siphunculi; BL 2.3-4.0 mm. Alatae without dark dorsal
abdominal cross-bands. On bark of twigs of numerous Populus and Salix
spp., common and widely distributed through North America, and in Mexico.
Oviparae and alate males in October-November (Palmer, 1952; Richards,
1967).

Pterocomma tremulae Börner Apterae dark warm brown or olive-brown,

wax-dusted, with straw-coloured siphunculi; BL 2.5-4.3 mm. Alatae have
variably developed dark dorsal abdominal sclerites, usually divided into
paired patches and not forming complete cross-bands and secondary rhinaria
distributed III 45-54, IV 0-7. Usually on suckers and two-year-old twigs of
Populus tremula, but occasionally found on other Populus spp. (see Heie,
1986). Attended by ants. In northwest, Central and Eastern Europe. Sexual
morphs da not appear to have been recorded. 2n = 8.
862 Pyrolachnus

Pterocomma xerophilae Ivanovskaya Apterae greenish-yellow to brownish;

BL c. 2.1 mm. On twigs of Salix xerophila and Salix sp in Western Siberia.
Monoecious holocyclic; alate male in late September (Ostanin, 1976). This
species could not be included in the key to Salix-feeders on the basis of
the available description. Apterae have well-developed wart-like marginal
tubercles (Ivanovskaya, 1977).

Pterocomma yezoense (Hori) Apterae orange-red to reddish-brown, with

antennae and legs mostly dark and siphunculi pale yellow with dark apices;
BL 2.9-3.8 mm. Alatae have dark dorsal abdominal cross-bands, sometimes
broken in the midline. On suckers, young stems and branches of Populus spp.
in Japan, maritime Siberia (Pashchenko, 1988b) and China (BMNH colln, leg.
VFE). Monoecious holocyclic; oviparae and alate males in November (Hori,
1929). Several other species described from Populus in China (anyangense,
neimongolense, sanpunum, sinipopulifoliae) are very similar (Zhang and
Zhong, 1980b), and their separate identity needs further confirmation.
2n = 8*.

PYROLACHNUS Basu and Hille Ris Lambers

Lachninae: Lachnini

Two or three species of very large East Asian aphids living on woody
Rosaceae. The alatae have an elongate pterostigma as in Longistigma, but it
does not curve around the tip of the wing. A.K. Ghosh (1982b) reviewed the
Indian species.

Pyrolachnus imbricatus David, Narayan and Rajasingh Apterae dull brown

with transverse rows of dark brown spots on dorsum, blackish tibiae and tarsi;
BL 4.4-5.2 mm. Alatae have only 9-15 secondary rhinaria on ANT HI (David
et al., 1971b). Feeding on stem and branches of Prunus spp., including
P. cornuta (Medda et al., 1986). In India (Himachal Pradesh, Uttar Pradesh).

Pyrolachnus pyri (Buckton) Apterae dull yellow to dark brown, often

dusted with wax meal, antennae and legs blackish; BL 3.5-6.0 mm. Alatae
have wings dark at their bases and secondary rhinaria distributed III 11-53.
IV 2-11, V 0-3 (variation perhaps reflecting degree of alatiformity). On
bark of branches of Pyrus communis and also recorded from Malus pumila
and Eriobotrya japonlca. In Iran, Bahrain, Pakistan, India, Nepal, Sri
Lanka, Korea and China. Holocyclic in China, where regular spring and
autumn migrations suggesting host alternation were observed (Long and
Chen, 1988). Apparently anholocyclic elsewhere. Pyrolachnus macrocanus
Zhang, described from Eriobotrya in China (Zhang and Zhong, 1982d), is
closely related and possibly a synonym.
 Radisectaphis 863

QUADRATUS Monzen Hormaphidinae: Nipponaphidini

One species forming galls on Distylium in Japan, having 4-segmented anten-

nae in all known morphs.

Quadratus yoshinomayai Monzen Galls on twigs of Distylium racemosum

are closed, pouch-like, max. diam. 2-4 cm, with numerous wart-like pro-
tuberances; greenish, becoming brown when mature, thick-walled, clothed
with short hairs. Alatae have 4-segmented antennae with secondary rhinaria
distributed III 22-27, IV 3-5, hyaline wings, yellowish-green abdomen, and
BL 1.6-2.0 mm. They emerge in early April from holes at apices of pro-
tuberances, flying to an unknown secondary host (Monzen, 1954; Takahashi,
1962a). Sorin (1987b) illustrated an embryo from an emigrant alata.

QUERNAPHIS Takahashi Hormaphidinae: Nipponaphidini

One oak-feeding species with small, flattened aptera having an oddly irregular
shape. Alatae have an unbranched media in the forewing and a hind wing
without any oblique veins.

Quernaphis tuberculata (Takahashi) Apterae yellow to white, not strongly

sclerotized, very variable in shape with legs concealed beneath body; BL
c. 0.5 mm. On undersides of leaves of Quercus gilva, feeding along the veins,
in Taiwan and Japan (Takahashi, 1958b). Alatae have dusky forewings, a pale
greenish-yellow abdomen and 4- or 5-segmented antennae with rhinaria
distributed III 9-10, IV 2-4, V (if present) 2-3 (Takahashi, 1933b). Apparently
entirely anholocyclic on Quercus, in both Taiwan and Japan.

RADISECTAPHIS Zhang Aphidinae: Pterocommatini?

Apterae and alatae described from Tibet (Zhang and Zhong, 198Ib) under the
name Radisectaphis gyirongensis appear to belong to a species of Pterocom-
matini. The recorded host plant is Malus pumila; presumably either this was
an error, or the aphids were vagrants.
864 Rectinasus

RECTINASUS Theobald Pemphiginae: Fordini

Probably only one species, with distinctive antennal sensoriation in alatae and
a long R IV+V. Affinities with other genera of Fordini were discussed by
Remaudière and Davatchi (1956).

Rectinasus buxtoni Theobald Galls on Pistacia spp. (palaestina, khinjuk)

are spindle-shaped sacs, 3-5 cm long and red in colour when mature, at or near
the base of the main vein of a leaflet (Fig. 126M). Alatae (BL 2.9-2.5 mm)
are produced in the third generation and leave the gall in September through
small longitudinal slits in the basal half. They start colonies on roots of
Compositae (Artemisia, Centaurea, Chondrilla, Sisymbrium) and also on
Gossypium and Papaver (Zumreoglu and Akbulut, 1984) and Gramineae
(BMNH colln). Roberti (1939) redescribed the apterous exules and sexuparae,
which have a very long R IV+V bearing numerous hairs. Holocyclic in
southwest Asia (Israel, Lebanon, Iran, Turkey), and populations on secondary
hosts are recorded from southern Europe (Italy, former Yugoslavia), North
Africa (Algeria), Kazakhstan, northern Caucasus, Transcaucasia, Dagestan
and Turkmenia. 2n = 26.

RETICULAPHIS Takahashi Hormaphidinae: Nipponaphidini

Three East Asian species having aleyrodiform apterae with heavily sclerotized
and reticulate dorsal cuticle. On Ficus or Fagaceae. At least one species forms
galls on Distylium, but the taxonomy and life cycle needs further investigation.
A.K. Ghosh and Raychaudhuri (1973b) reviewed the genus.

Reticulaphis distylii (van der Goot) Galls on Distylium stellare in Java are
conical or fingertip-like, brownish-green when mature (Hille Ris Lambers and
Takahashi, 1959). The gall illustrated (as Schizoneuraphis distylii van der
Goot) by Docters van Leeuwen-Reijnvaan and Docters van Leeuwen (1926)
seems to be another species (see Noordam, 1991, p. 259). Alatae (BL 1.5-
1.9mm), secondary rhinaria distributed III 19-31, IV 8-12, V 6-10, abdomen
dark green) are present in the galls in October. Host alternation apparently
occurs to Ficus spp. (Hille Ris Lambers and Takahashi, 1959), but this needs
to be confirmed experimentally; very similar aphids, identified as subspecies
rotifera, have been collected from Quercus sp. in India (Raychaudhuri et al.,
1980c). Apterae on Ficus are dark brown to dull purplish-black, acquiring with
age a fringe of wax; BL 0.5-0.9 mm. Immature apterae are yellow-brown and
active. Immature alatae on Ficus have a woolly wax fringe plus long erect wax
 Rhopalosiphoninus 865

filaments; when adult they are dark violet-black, without wax. Zimmerman
(1948) provided a well-illustrated account of this species in Hawaii. Hille Ris
Lambers and Takahashi (1959) recognized several 'subspecies' on Ficus in
Taiwan and Java and provided a key, noting that many of the differences
could prove to be within the range of variation of one species. The most
commonly encountered form on Ficus, R. distylii fici (Takahashi), has
marginal hairs about 75 µm long with acute apices, and is recorded from
China, Taiwan, Japan, Java, Nepal (BMNH colln, leg. K.C. Sharma) and the
Philippines. Another fairly distinct form, R. distylii foveolatae, is smaller and
narrower (BL c. 0.5 mm) and has no evident wax secretion; it appears specific
to Ficus foveolata in mountainous regions of Taiwan and Japan (Takahashi
1935c, 1958a).

Reticulaphis mirabilis (Takahashi) Apterae oval, black, strongly sclerotized,

without marginal wax; BL c. 0.4mm. Attached firmly to undersides of leaves
of Ficus sp. in Taiwan (Takahashi, 1939a), and also recorded from Thailand
(Tao, 1966). Other morphs and life cycle unknown.

Reticulaphis shiiae Takahashi Apterae oval, black, with black legs; BL

0.7-0.9 mm. On leaves of Castanopsis cuspidata in Japan (Takahashi, 1958a).
Other morphs and life cycle unknown.

RHOPALOSIPHONINUS Baker Aphidinae: Macrosiphini

About 20 species with large and strongly swollen siphunculi and a short
triangular CAUDA, living on a great variety of plants, often in cryptic
habitats. Three species are recorded from trees; at least two of these have host
alternation to herbaceous plants. See also B & E, 1984, p. 337.

Rhopalosiphoninus ehretis Bhattacharya and Chakrabarti Colour in life not

recorded, probably apterae are pale; BL 1.9-2.1 mm. Forming spiral leaf-rolls
in spring on Ehretia acuminata in Uttar Pradesh, India (Bhattacharya and
Chakrabarti, 1982). Alatae have a brown dorsal abdominal patch. Life cycle
unknown.

Rhopalosiphoninus staphyleae (Koch) Apterae on Staphylea in spring are

yellowish-white or pale yellow with a translucent whitish spot on anterior part
of dorsal abdomen; BL 2.3-3.0mm. Causing leaves to curl and become
mottled pale yellow. Heteroecious holocyclic; alatae, with a large black dorsal
abdominal patch, migrate from late May onwards to found colonies par-
ticularly on Liliaceae and Iridaceae, but also sometimes on plants in at least
11 other families, including a few records from trees (e.g. Forbes and Chan,
1989). Apterae on secondary hosts, and also those in populations remaining
866 Rhopalosiphum

through summer on Staphylea, are much more pigmented than the spring
generations; green, with variably-developed olive green to black dorsal
abdominal cross-bands. The holocycle is only recorded from Europe, but
anholocyclic populations of staphyleae group are almost cosmopolitan (see
B & E, 1984, p. 339). 2n = 10.

Rhopalosiphoninus tiliae (Matsumura) Apterae on Tilia in spring are

yellowish-green with brownish antennae and mainly black legs and siphun-
culi; BL 2.7-3.0 mm. On undersides of young leaves, which become curled
(Miyazaki, 1971). Heteroecious holocyclic; alatae with a large black dorsal
abdominal patch migrate from early June onwards to found colonies on
leaves and flower-stems of Adenocaulon himalaicum (Miyazaki, 1985). In
Japan and eastern Russia (Pashchenko, 1984). 2n = 12 (Shinji, 1941, as
nobukii).

RHOPALOSIPHUM Koch Aphidinae: Aphidini

About 13 species associated with Prunus or Pyroidea as primary hosts, with

Gramineae, Cyperaceae or more rarely other plants as secondary hosts. There
are close affinities to Melanaphis and Schizaphis. Accounts are available
for Britain (Stroyan, 1984), northwest Europe (Heie, 1986), northeast India
(Raychaudhuri, 1980), Japan (Takahashi, 1965b), Australia (Eastop, 1966),
Canada (Richards, 1960) and North America (Richards, 1962). See also
B & E, 1984, pp.339-343.

Rhopalosiphum arundinariae (Tissot) Apterae yellowish-brown with darker

brown head and sides of body; BL c. 1.6mm. Dense colonies were found
in April on Arundinaria tecta in Florida, USA (Tissot, 1933). Life cycle
unknown.

Rhopalosiphum cerasifoliae (Fitch) Apterae in spring colonies are pale

yellowish to pale green with spinal, marginal and intersegmental lines of
darker green pigment and a dusting of wax; BL 2.0-2.5 mm. On leaves and
shoots of Prunus virginiana and P. pennsylvanica, distorting growing tips
and rolling the leaves. Heteroecious holocyclic; alatae migrate in May-June
to Scirpus and Eleocharis (Voegtlin and Halbert, 1990), but populations some-
times also persist on the primary host (Richards, 1960). Oviparae on Prunus
in autumn are yellow with siphunculi pale-dusky (cf. R. padi). Widely dis-
tributed in North America. 2n = 8.

Rhopalosiphum insertum (Walker) Apterae in spring colonies are rather

shiny, bright green to yellow-green with a dark green spinal stripe; BL
2.1-2.6 mm. Curling young leaves of Pomoidea (Cotoneaster, Crataegus,
 Rhopalosiphum 867

Malus, Pyrus, Sorbus and sometimes on Cydonia and Mespilus). It may also
occasionally form colonies on suckers of Prunus cerasus. Heteroecious
holocyclic; alatae fly in late May-June to found colonies on subterranean
parts of various Gramineae, but colonies may persist into summer on primary
hosts. In Europe, eastward to the Urals and Turkey; the Azores; possibly
originating from North America (where for many years the name fitchii was
used for insert m on its primary hosts). Evenhuis (1968) studied its natural
control and the role of a parasitoid in apple orchards. See also B & E, 1984,
p. 340. 2n = 10.

Rhopalosiphum maidis (Fitch) The holocycle of this well-known cereal pest

(see B & E, 1984, p. 340) was recently found to occur in Pakistan, with Prunus
cornuta as primary host. Fundatrices and their wax-powdered apterous pro-
geny feed along veins on undersides of deformed leaves, in mixed colonies with
Myzus cornutus (Remaudière and Naumann-Etienne, 1991). Alatae migrate to
Gramineae before the end of May. The primary host morphs have not yet
been fully described; although Aphis cooki Essig, described from Citrus
in California (Essig, 1911), could be based on the second generation of
Rh. maidis. 2n = 8, 9 or 10.

Rhopalosiphum musae (Schouteden) (= scirpifolii Gillette and Palmer)

Apparently heteroecious between Prunus and marsh or water plants (e.g.
Scirpus, Typha) in US Rocky Mountain region from Washington to Colorado.
Oviparae and alate males were described from Prunus besseyi in Colorado
in October-November (Gillette and Palmer, 1932; as scirpifolii), and some
sexuales collected along with gynoparae on P. subcordata in Oregon in
October (BMNH colln, leg. L.G. Gentner) agree well with that description.
However, the spring generations have not been described, so we are not able
to include this species in the key to Prunus aphids. Rhopalosiphum musae was
originally described from a glasshouse population on Musa ensete in Belgium;
specimens from presumably anholocyclic populations on secondary hosts in
Cyperaceae and Araceae, or trapped alatae identified as this species, are
recorded from England, Germany, Iran, Rhodesia, Tanzania and Australia
(BMNH colln).

Rhopalosiphum nigrum Richards Apterae in spring colonies on Crataegus

sp(p). are dark green to blue-black, usually dusted with wax; BL 1.7-2.2 mm.
Heteroecious holocyclic; alatae migrate in late May-early July to grasses and
sedges in damp situations (Richards, 1960). In Canada (Ontario, Manitoba)
and USA (Oregon, Utah).

Rhopalosiphum nymphaeae (Linnaeus) Apterae in spring colonies reddish-

brown to dark olive, dusted with light grey wax especially on ABD TERG 1-4;
BL 1.6-2.6 mm. Feeding on young twigs, leaf petioles and fruit stalks of
various Prunus spp., attended by ants. Heteroecious holocyclic; alatae fly in
May-June to a variety of water plants. Virtually worldwide. See also B & E,
1984, p. 341. 2n = 8.
868 Roepkea

Rhopalosiphum padí (Linnaeus) Apterae in spring colonies on Prunus are

pale green to dark green, brown or nearly black, with a rust-red patch
around base of each siphunculus, and a coating of mealy wax; BL 2.0-2.8 mm.
Heteroecious holocyclic; the primary host is usually Prunus padus in Europe,
feeding on undersides of young leaves which become longitudinally rolled or
folded to enclose the aphid colony. In North America the main primary host
is P. virginiana and the leaf distortion may be less (Richards, 1960). Alatae
migrate in May-June to numerous grasses and cereals (see B & E, 1984,
pp. 341-342), but colonies may also persist into summer on Prunus. Distribu-
tion is now virtually cosmopolitan, anholocyclic populations occurring in
warm climates or where primary hosts are unavailable. Leather (1988)
reviewed its biology on P. padus in Europe, Loxdale and Brookes (1988) and
Sherlock et al. (1986) respectively studied enzyme variation and elemental
composition in holocyclic populations, and Tatchell and Parker (1990) studied
selection of the primary host by return migrants. 2n = 8 (or sometimes 9;
Hales and Cowan, 1990).

Rhopalosiphum rufiabdominalis (Sasaki) Apterae in spring colonies on

young leaves, stems and suckers of Prunus spp. (mume, yedoensis) in East
Asia are reddish or greenish-brown with bluish-white mealy wax on sides of
body and forming dorsal cross-bands (Moritsu, 1983); BL 2.0-2.6 mm.
Heteroecious holocyclic (Tanaka, 1961); alatae migrate in May-June to form
colonies on underground parts of numerous species of Gramineae, Cyperaceae
and some dicots. Anholocyclic populations occur throughout most of the
world on secondary hosts. See also B & E, 1984, p. 343. 2n = 8.

Rhopalosiphum rufulum Richards Apterae in spring colonies on Crataegus

in Canada are mainly maroon, mottled with green or yellow, with a pattern
of mealy wax; BL 2.0-2.5 mm (Richards, 1960). Presumably heteroecious
holocyclic, alatae migrating in May-June; however, although collected from
Crataegus sp(p). in spring in Manitoba, Quebec, Ontario and New Brunswick,
secondary host populations have only been found in Europe (former Czecho-
slovakia, Denmark, England, Germany, The Netherlands), where large
populations have occurred since about 1970 on Acorus calamus (see Heie,
1986). Stroyan (1972) reared alate gynoparae and males in England and
obtained oviparae, eggs and fundatrices on Crataegus monogyna. 2n = 8.

ROEPKEA Hille Ris Lambers Aphidinae: Macrosiphini

One (or two) palaearctic species, resembling Nearctaphis but without lateral
abdominal tubercles, and the primary host is Prunus rather than Pomoidea.
Roepkea marchali (Börner) Apterae a dirty yellowish-green to almost black,
according to degree of dorsal sclerotization; BL 2.0-2.3 mm. Alatae have
 Sanbornia 869

a black dorsal abdominal patch. Feeding in spring on undersides of leaves

of Prunus mahaleb, which become rolled into broad tubes, inflated and
yellowed. In southern Europe (France, Italy) alatae are rarely produced and
there is apparently no host alternation, but East European and southwest
Asian populations, regarded as a separate subspecies, R. marchali bathi-
aschvili Abashidze, produce more numerous alatae in July-August and seem
to have at least a partial host alternation to flowers of Labiatae (Stachys,
Phlomis). This eastern form, which is recorded from Crimea, Georgia, Israel,
Iran, Lebanon and Turkey, has a shorter antennal PT and the apterae on
Prunus generally have dark dorsal cross-bands rather than a completely
sclerotic tergum (Hille Ris Lambers, 1966a). Experimental confirmation of
the life cycle is needed. 2n = 12.

RYOICHITAKAHASHIA Hille Ris Lambers

Aphidinae: Aphidini

One species in Japan, resembling Aphis in its general appearance and arrange-
ment of marginal tubercles, but with numerous marginal hairs on abdominal
segments; with somewhat developed antennal tubercles, giving the front of the
head a concave outline in dorsal view; and with secondary rhinaria on ANT
III (and sometimes IV) in apterae.

Ryoichitakahashia prunifoltae (Shinji) Apterae broad-bodied, rather dull

reddish-black with banded antennae, central sections of tibiae pale and jet
black siphunculi (see Moritsu, 1983; as Macchiatiella ilexis); BL 2.1-2.2 mm.
In dense colonies on stems of Ilex serrata (var. sieboldii) in Japan. Alatae have
a black dorsal abdominal patch. The nomenclature is confused; Hille Ris
Lambers (1965) pointed out that this aphid was illustrated by Shinji (1941)
under the name Anuraphis celastri. Life cycle unstudied; oviparae were
collected on Ilex in early November (BMNH colln, leg. M. Sorin), but it is
not known whether host alternation occurs.

SANBORNIA Baker Aphidinae: Aphidini(?)

One North American species on Juniperus with uniquely specialized mor-

phology and therefore of uncertain phylogenetic position, but perhaps closest
to Siphonatrophia, In addition to features mentioned in the key to juniper
aphids, marginal abdominal tubercles are absent and the reduced tarsi have
hairless first segments.
870 Sappaphis

Sanbornia juniperi Pergande ex Baker, Plate 13e, f Apterae green, with a

dorsal coat of flaky, transparent wax (Boudreaux, 1949); BL 1.2-1.4 mm.
Alatae have pale brownish-green head and thorax. On Juniperus (scopularum,
virginiana) in USA. Monoecious holocyclic; oviparae and alate males in
October-November. Trioxys cupressicola may be a specialized parasitoid.

SAPPAPHIS Matsumura Aphidinae: Macrosiphini

A small oriental genus with Pyrus as primary hosts, related to Dysaphis but
with long hairs on body and antennae. Accounts are available for Japan
(Miyazaki, 1971) and China (Zhang and Zhong, 1983b). The name Sappaphis
was used in a broader sense from 1952 to 1965 to include species now in
Dysaphis.

Sappaphis piri Matsumura Apterae in spring on Pyrus are broadly oval,

brownish or yellowish-brown, covered with white woolly wax; BL 1.8-2.5 mm.
Alatae have dark dorsal abdominal cross-bars, more weakly developed on
anterior tergites; secondary rhinaria distributed III 21-28, IV 4-9, V 0-1. On
undersides of leaves of Pyrus spp. in East Asia (Japan, Korea, China, Taiwan,
eastern Siberia). Heteroecious holocyclic, migrating in May-June to roots of
Artemisia spp. (where it is orange-red in colour and without wax); returning
to pear in October. Kim et al. (1986) studied seasonal population fluctuations
on pears in Korea, and Takada and Hashimoto (1985) studied relations with
parasitoids and ants on the secondary host. 2n = 12. [Zhang (in Zhang and
Zhong, 1980a) described two species of Sappaphis from spring populations on
unidentified Pyrus in China. Sappaphis dipirivora Zhang has ANT PT much
shorter than base in both apterae and alatae, and 2-5 secondary rhinaria on
ANT III in apterae; S. sinipiricola Zhang has short siphunculi with 3-4 hairs,
and many more secondary rhinaria in alatae (distributed III 39-45, IV 11-15,
V 0-9). Life cycle of both species unknown. Sappaphis sinipiricola has
2n = 12 (Chen and Zhang, 1985b).]

SCHIZAPHIS Börner Aphidinae: Aphidini

About 36 palaearctic and six nearctic species resembling Rhopalosiphum but

with tapering siphunculi and media of forewing only once branched. Most live
all year on Gramineae or Cyperaceae but a few mainly Asian species over-
winter as eggs on Pyrus. The taxonomy and life cycles of the pear-feeding
species require further study. Eastop (1961) provided a key to the world fauna
then known. Pear-feeding species were keyed by Shaposhnikov (1952), and
 Schizolachnus 871

included in regional accounts for northeast India (Raychaudhuri, 1980);

eastern Siberia (Pashchenko, 1988b) and China (Tao, 1962). See also B & E,
1984, pp. 343-346.

Schizaphis piricola (Matsumura) Apterae yellowish-brown or green, with

dark antennae and black-tipped siphunculi; BL c. 1.4 mm. Spring colonies live
in rolled edges of leaves of Pyrus pyrifolia (Essig and Kuwana, 1918). In
Japan, Korea and China. Alatae are produced in May and presumably fly to
an unknown secondary host. 2n = 8 (Chen and Zhang, 1985b).

Schizaphis punjabipyri (Das) Apterae greyish or blackish, dusted with white

wax, with black siphunculi and CAUDA; BL c. 1.9 mm. Feeding by fundatrices
on developing leaves of Pyrus communis in early spring prevents them from
unfolding and causes the edges of the leaves to stick together forming a slightly
inflated pod-shaped gall. Recorded from northern India, Pakistan and Nepal.
Alatae migrate in May to an unknown secondary host. In Pakistan gynoparae
and alate males return to Pyrus in early December or earlier, oviparae laying
eggs in late December-January which hatch in February-March (Das, 1918).

Schizaphis pyri Shaposhnikov Apterae green with dark siphunculi and

CAUDA; BL c. 1.7mm. In spring colonies on Pyrus communis, feeding on
upper sides of leaves and folding them longitudinally in half, with the underside
turned outward, to form a similar pod-shaped gall to that of S. punjabipyri
(Shaposhnikov, 1952). Apparently heteroecious holocyclic, alatae migrating in
May to form colonies on Cyperaceae (Leclant, 1967; as S. fritzmuelleri), but
this needs experimental confirmation. Recorded from southern France (on
Carex); Azores, Madeira, Canaries (Nieto Nafria, 1985); European Russia;
Kazakhstan; and eastern Siberia (Pashchenko, 1988b). Sexual morphs have
not been described.

SCHIZOLACHNUS Mordvilko Lachninae: Cinarini

Seven or eight species of oval-bodied, hairy, wax-secreting aphids, all feeding

on Pinus needles; four or five nearctic, two western palaearctic and one
eastern palaearctic. Phylogenetic relations with Cinara and Eulachnus were
studied by Sorensen (1990), who concluded that Schizolachnus was more
closely related to Eulachnus. Hottes (1956a) keyed the North American species
then recognized. Pauesia unilachni is a specialized parasitoid.

Schizolachnus curvispinosus Hottes, Essig and Knowlton Colour of aptera

in life not recorded, 'probably a dusky green', with black hind tibiae; BL
2.1-2.8 mm. On needles of Pinus ponderosa, and occasionally found on
872 Schizolachnus

non-native pines, in western North America. Oviparae and alate males occur
in October (Hottes et al., 1954). The alata was described by Hottes (1956d).

Schizolachnus flocculosus (Williams) Apterae have black head and thorax

with grey wax markings, abdomen light brown or green with narrow cross-
bands of wax (Hottes, 1952a; as S. wahlea) and black hind legs; BL 2.4-
2.9mm. On needles of Pinus arizonica and P. ponderosa in US Rocky
Mountain region (Colorado, Utah) and Arizona. Oviparae and alate males in
September-November (Hottes, 1954b; as S. wahlea).

Schizolachnus orientalis (Takahashi) Apterae yellowish-brown in original

description from Taiwan (Takahashi, 1924a), but recorded as blackish-brown
in India (A.K. Ghosh, 1982b), covered in fine, dirty white wax meal; BL
1.9-2.4 mm. Feeding in rows along needles of Pinus spp. in India, Japan,
Korea, China and Taiwan. Oviparae and alate males in October-November
(Inouye, 1970).

Schizolachnus parvus (Wilson) Apterae brownish tinged with green, covered

with bluish-white wax threads which also extend onto needles; BL 1.6-1.8 mm.
On needles of Pinus spp. in eastern USA. Oviparae and alate males in late
October (Wilson, 1915). Schizolachnus lanosus Hottes (1959) appears to be
a synonym.

Schizolachnus pineti (Fabricius), Plate 16e Apterae dark greyish-green

covered in wax meal giving a light bluish-grey appearance; BL 1.2-2.5 mm.
On numerous Pinus spp., especially on young trees, forming dense colonies
in rows along the previous year's needles. Oviparae and alate males in
October-December; in some years colonies persist into early winter and
possibly anholocyclic overwintering occurs in mild winters (Carter and
Maslen, 1982). Common and widespread in Europe; also in Kazakhstan,
Siberia (Pashchenko, 1988b), China (Zhang and Zhong, 1985e) and intro-
duced to North America. Holman (1990) studied morphological changes in
spring generations, Kidd et al. (1985) studied the association with Eulachnus
agilis and Thompson (1977) studied its effect on tree growth. Some popula-
tions on Pinus nigra (also on mugo, pinea; BMNH colln) in Europe, and
east to Turkey, have a long tip to the rostrum and more hairs on ANT II,
corresponding to S. pineti ssp. obscurus Börner, which should perhaps be
regarded as a distinct species (Klimaszewski and Wojcieckowski, 1976).
2n = 18 (Russia; Rukavishnikov, 1972) or 2n = 10 (England; Blackman,
1980); again indicating that there is more than one species under this name.

Schizolachnus piniradiatae (Davidson), Plate 16f Apterae dark olive green

to brown, with grey flocculent wax coating; BL 2.0-2.5 mm. On needles of
numerous Pinus spp., widely distributed in North America. Oviparae and
alate males occur in September-November. There have been studies in Canada
of its life history and ecology (Gröbler, 1962), population dynamics on Pinus
resinosa (Sharma and Laviolette, 1968a), oviposition behaviour (Sharma and
 Schizoneuraphis 873

Laviolette, 1968b), predators (Gagné and Martin, 1968) and fungal diseases
(Gröbler et al., 1962; Tyrrell and Macleod, 1975).

SCHIZONEURAPHIS van der Goot

Hormaphidinae: Nipponaphidini

Six species are known, at least one of them having host alternation between
Distylium galls and Litsea (in Java; Noordam, 1991). Closely related to
Nipponaphis, but the aleyrodiform exules have long, basally thick dorsal
(prosomal) hairs.

Schizoneuraphis gallarum van der Goot Galls on Distylium stellare in Java

are oval, balloon-shaped, 25-50 mm long, apparently arising from the stem
but formed from the leaf (Fig. 130J, p. 785); pale green to peach-coloured
when young, becoming dark grey-brown and woody, but with a thin circular
window in the side which becomes the emergence hole for alatae in August-
October. Emigrant alatae (BL 1.6-2.2 mm) have secondary rhinaria distributed
III 17-22, IV 8-11, V 4-8. Apterous exules feeding along veins on undersides
of leaves of Litsea chinensis are aleyrodiform, shiny dark brown to black, with
upright pale brown dorsal hairs and a white marginal band of wax; BL
0.7-0.9 mm. Host alternation still needs to be confirmed experimentally. Hille
Ris Lambers and Takahashi (1959) claimed to have established it on the basis
of similarity of embryos within emigrant alatae with first instars on Litsea,
but Noordam (1991) noted some differences.

Schizoneuraphis himalayensis (Ghosh and Raychaudhuri) Apterae broadly

oval, dark brown to black; BL 0.9-1.1 mm (A.K. Ghosh and Raychaudhuri,
1973b). In colonies on leaves and stems of Litsea polyantha and Persea
bombycana in India (Assam, West Bengal). Alatae (sexuparae?) were pro-
duced in January. Life cycle unknown. Nipponaphis machiliphaga in A.K.
Ghosh et al. (197Id) appears to be this species.

Schizoneuraphis litseicola Noordam Apterae on undersides of leaves of

Litsea amara are aleyrodiform, shiny brown-black, similar in life to S.
gallarum; BL 0.8-0.9 mm. Only known from Java (Noordam, 1991). Life
cycle unknown.

Schizoneuraphis longisetosus Noordam Galls on Distylium stellare in Java

are not properly described, but believed to be semiglobular, turning red when
mature (possibly gall No. 487 of Docters van Leeuwen-Reijnvaan and Docters
van Leeuwen, 1926). Emigrant alatae (BL 2.4-2.8 mm, secondary rhinaria
distributed III 25-31, IV 12-16, V 8-13) were collected in late May (Noordam,
1991). Secondary host populations not known.
874 Schizoneurata

Schizoneuraphis machiliphaga (Takahashi) Apterae almost circular, dor-

sally flattened, reddish-brown, not strongly sclerotized; BL 0.8-1.0 mm. On
undersides of leaves of Persea sp. in Japan (Takahashi, 1959b). Life cycle
unknown.

Schizoneuraphis malayna (Takahashi) Apterae aleyrodiform, almost cir-

cular, dorsally flattened, shiny brownish-black, without evident wax; BL
c. 1.3mm. On undersides of leaves of Quercus sp. in Cameron Highlands
of Malaya (Takahashi, 1950; as Thoracaphis malaynus). Life cycle unknown.

Schizoneuraphis querciphaga (Ghosh and Raychaudhuri) Apterae yellowish

to dark brown; BL 1.3-1.5 mm. On apical stems of Quercus sp., attended by
ants, in Sikkim, India (A.K. Ghosh and Raychaudhuri, 1973b), and also on
Castanopsis in Nepal (Miyazaki, 1977). Life cycle unknown. Very similar to
S. malayna.

SCHIZONEURATA Hille Ris Lambers

Pemphiginae: Eriosomatini

One little-known North American species closely related to Eriosoma but

lacking siphunculi in morphs developing on Crataegus, the presumed secon-
dary host. The primary host is unknown; presumed to be Ulmaceae (Hille Ris
Lambers, 1973c).

Schizoneurata tissoti Hille Ris Lambers Only immatures and alate sexuparae
are described, and appearance in life is unknown. Apterae and immatures
are presumably wax-covered. Sexuparae have BL 1.6-2.2 mm and secondary
rhinaria distributed III 15-20, IV 2-5, V 1-3, VI 0-1. On Crataegus uniflora
and Crataegus sp. in Florida, USA (Hille Ris Lambers, 1973c).

SCHIZONEURELLA Hille Ris Lambers

Pemphiginae: Eriosomatini

One species in the Himalayan region, with empodial hairs reduced (in
alatae) or absent (in apterous exules); related to Eriosoma but without
siphunculi in the morphs on the secondary host plant, and distinct from
the nearctic genus Schizoneurata because it has embryos with spinulose
hind tarsi, as in palaearctic Eriosoma (Schizoneura) (Hille Ris Lambers,
1973b).
 Schlechtendalia 875

Schzoneurella indica Hille Ris Lambers Galls are clavate, asymmetric

pouches arising from leaves of Ulmus spp. (laevigata = villosa, wallichiana)
in India (Kashmir, Jammu) and Pakistan (Hille Ris Lambers, 1973b; Bhagat,
1982b). Alatae (BL 1.8-2.3 mm) are produced in May-June. Host alternation
occurs to Malus pumila, where colonies (presumably wax-covered) live on the
roots. Bhagat (1982b) reported a parasitoid and hyperparasitoid of the gall-
feeding generations.

SCHLECHTENDALIA Lichtenstein Pemphiginae: Fordini

Two or three East Asian species related to Melaphis, but alatae emerging from
galls on Rhus have an elongate, curved pterostigma.

Schlechtendalia chinensis (Bell), Plate 6f Galls ('Chinese gall-nuts') are

greenish-yellow, densely covered with short, fine hairs, ovoid, becoming
woody and irregular in shape with horn-like protrusions, varying from 1 to
6 cm in maximum length (Fig. 133). On Rhus semialata ( = javanica) in China,
Japan, Korea, Taiwan and Malaya. Heteroecious holocyclic, with an unusual
one-year cycle; after several generations in the gall, emigrant alatae (BL
c. 1.5mm) are produced in October and fly to found overwintering colonies
on mosses (Mnium, Plagiomnium). Alatae sexuparae return to Rhus in spring
to produce sexuales. Oviparae live in bark crevices for three weeks after
mating and then each produces a single fundatrix larva ovoviviparously - a
phenomenon not known in any other aphid (Tang, 1976; Takada, 1991a). The
galls have been collected for many years; they have been used in tanning
leather, in the preparation of dyes and for medicinal purposes (including a

Fig. 133. Gall of Schlechtendalia chinensis on Rhus semialata (after Baker, 1917b).
876 Schoutedenia

treatment for haemorrhoids; Gardener, 1981). Takada (1991b) studied gall

development. 2n = c. 36.

Schlechtendalia elongallis (Tsai and Tang) Galls are elliptical, maximally

4.1 cm long, green, becoming red or purplish when mature with many fine
longitudinal ridges. On undersides of leaflets of Rhus punjabensis var. sinica
in China (Tsai and Tang, 1946). Alatae (BL c. 1.3 mm) emerge through a large
slit in autumn. Life cycle unknown.

Schlechtendalia microgallis (Xiang) Galls are small, papilliform, on Rhus

potanini in China (Xiang, 1980/81; as Meitanaphis microgallis). Closely
related to S. elongallis and possibly a synonym.

SCHOUTEDENIA Rübsaamen Greenideinae: Cervaphidini

Two species on woody Euphorbiaceae belonging to an ancient group (also

including Eonaphis and Paulianaphis) of Gondwanaian distribution.
Remaudière (1988, 1990) reviewed the genus, synonymizing most of the
described species.

Schoutedenia emblica Patel and Kulkarni Apterae yellow-green to green;

BL 1.4-1.6 mm. In ant-attended colonies, only on Phyllanthus emblica
(= Emblica officinalis) in India, Nepal and Thailand. Monoecious holocyclic;
in southern India, apterous (brachypterous?) males and oviparae are produced
in August (David and Hille Ris Lambers, 1956; as ssp. andrhaka).

Schoutedenia ralumensis Rübsaamen (= S. lutea), Plate l1e, f Apterae

lemon-yellow or green; BL 1.4-1.8 mm. Alatae have thickly black-bordered
forewing veins. On young shoots, leaves and stems of woody Euphorbiaceae
(Breynia, Bridelia, Fluggea, Glochidion, Phyllanthus), attended by ants.
Widespread in Southeast Asia; India; Africa; and down the east coast of
Australia. Monoecious and partially holocyclic; oviparae and alate males
occur concurrently with parthenogenetic generations throughout summer and
autumn. Presumably all eggs laid enter diapause, as fundatrices occur only
in spring (Hales and Carver, 1976). The oviparae are unusual in having scent
glands ('pseudosensoria') on the hind femora - although in S. emblica they are
on the hind tibiae as in other aphids. Relations with predators and parasitoids
in Australia were studied by Hales and Carver (1976). 2n = 16 (female),
14 (male) or 15* (sex unknown; Papua New Guinea, leg. J.H. Martin);
the highly anomalous spermatogenesis of this aphid was studied by Hales
(1989).
 Shivaphis 877

SERRATOCALLIS Quednau and Chakrabarti

Drepanosiphinae: Phyllaphidini

One oak-feeding species in India, related to Hoplochaetaphis (Quednau and

Chakrabarti, 1976).

Serratocallis takahashii Quednau and Chakrabarti Apterae brown (D.C.

Gautam, pers. comm.); BL 1.6-2.3 mm. Alatae have a short broad ptero-
stigma with a black spot at its base. On stems and leaf petioles of Quercus
dealbata (BMNH colln, leg. D.C. Gautam), Q. baloot in Afghanistan and
Quercus sp. in India (Himachal Pradesh, Uttar Pradesh; A.K. Ghosh and
Quednau, 1990). Sexual morphs unknown.

SHENAWHEUM Hottes and Frison

Drepanosiphinae: Drepanosiphini

One North American species on Acer, related to Drepanaphis and considered

as a subgenus by some recent authors; the differences were discussed by Smith
and Dillery (1968).

Shenawheum minutum (Davis) All viviparae alate, with brown or black

thorax and very pale greenish-yellow to whitish abdomen; BL 1.0-1.2 mm.
Immatures are pale greenish-yellow. On undersides of leaves of Acer sac-
charum in eastern North America. Sexual morphs are apparently unrecorded.

SHIVAPHIS Das Drepanosiphinae: Phyllaphidini

Six Asiatic species mainly associated with Celtis, with one described from
Tilia. Quednau and Remaudière (1985) keyed the species then known, and
A.K. Ghosh and Quednau (1990) reviewed the Indian species.

Shivaphis catalpinari Quednau and Remaudière Appearance in life not

recorded, perhaps like Sh. celti; BL of aptera 1.4-1.7 mm, of alata 1.5-
1.8 mm. On Celtis australis in Turkey. Monoecious holocyclic; oviparae and
alate males in October (Quednau and Remaudière, 1985).
878 Siculaphis

Shivaphis celti Das, Plate l0c, d Apterae pale to dusky greenish, sometimes
pinkish, covered in white flocculent wax; BL 1.9-2.6 mm. Alatae also wax-
covered, with forewing veins thickly bordered with fuscous distally and
large black patches on pterostigma and distal part of Cu lb . On undersides
of leaves or on shoot tips of Celtis spp. in Turkey, India, Pakistan, Nepal,
Sri Lanka, Korea, Japan, Loochoo, China, Taiwan and Philippines. Mono-
ecious holocyclic; Das (1918) gave a full account of its biology and natural
enemies in Pakistan; oviparae and alate males occurred in December, eggs
were laid in January and hatched in March. In Japan, Shibata (1955; as
Sh. celticolens) obtained sexuales in September-November. In Taiwan,
Sh. celti is mainly anholocyclic, but produces some sexuales in December-
January. 2n = 10.

Shivaphis celticola (Nevsky) All viviparae alate, pale green to greenish-

yellow covered with bluish wax; BL 1.8-2.2 mm. Forewings are marked with
fuscous as in Sh. celti, except that there is a band joining the first fork of the
media and Cu la (A.K. Ghosh and Quednau, 1990). On undersides of leaves
of Celtis caucasica (= australis), recorded from Tadzhikistan, Uzbekistan
and India (Kashmir). Monoecious holocyclic, with oviparae and alate males
in October-November; Narzikulov (196la) gave a detailed account of the
life cycle in Tadzhikistan, with descriptions of all morphs.

Shivaphis (Sinishivaphis) hangzhouensis (Zhang and Zhong) Only alate

viviparae known; yellow, covered with wax, BL 1.9-2.3 mm. Described from
Osmanthus asiaticus (Zhang and Zhong, 1982c), but since found on under-
sides of leaves of Celtis julianae (BMNH colln, leg. VFE), which is a more
likely host. Life cycle unknown. 2n = 10*.

Shivaphis szelegiewiczi Quednau Only alate viviparae known; colour in life

not recorded, presumably covered in wax, BL c. 2.5mm. Forewings are
marked with fuscous as in Sh. celti, but with a spot below the pterostigma,
next to base of media. On Celtis aurantiaca in Korea (Quednau, 1979). Life
cycle unknown.

Shivaphis (Sinishivaphis) tilisucta Zhang Only alate viviparae known; bright

green covered with thick wax powder, BL c. 1.9mm. On undersides of leaves
of Tilia tuan in China (Zhang and Zhong, 1990). Life cycle unknown.

SICULAPHIS Quednau and Barbagallo

Drepanosiphinae: Phyllaphidini

One species in southern Italy on Quercus, the apterae of which have remark-
able variation in form of dorsal hairs.
 Sinolachnus 879

Siculaphis vittoriensis Quednau and Barbagallo Apterae pale yellowish-

green with longitudinal olive green dorsal stripes leaving a pale spinal line;
BL 1.0-1.5 mm. Alatae have smoky wings with Cu lb strongly bordered with
fuscous. On Quercus calliprinos, forming ant-attended colonies mainly along
mid-ribs on basal parts of undersides of leaves. Only known from Sicily.
No sexual morphs or fundatrices were found, and anholocycly is suspected
(Quednau and Barbagallo, 1991).

SIGMACALLIS Zhang Pemphiginae: Pemphigini

One species, Sigmacallis pilosa Zhang, is described from alatae collected on

Populus sp. in Tibet (Zhang and Zhong, 1981b). It appears to be a species
of Pachypappa, q.v.

SINOCHAITOPHORUS Takahashi
Drepanosiphinae: Phyllaphidini

Two species on Ulmus in East Asia related to Indochaitophorus. Quednau

and Shaposhnikov (1988) discussed the differences between the species, and
between this and related genera.

Sinochaitophorus maoi Takahashi Colour of aptera not recorded, probably

blackish-brown; BL c. 1.3 mm. Alatae blackish-brown, mottled with red, with
maculate wings (Holman and Szelegiewicz, 1974a). On Ulmus pumila in
Mongolia and China. Biology and life cycle unknown.

Sinochaitophorus mordvilkoi Quednau and Shaposhnikov Colour of aptera

in life not recorded, probably dark brown or blackish; BL c. 1.3mm.
Along mid-ribs on undersides of leaves of Ulmus japonica in eastern Russia
(Quednau and Shaposhnikov, 1988). Alatae and other morphs unknown.

SINOLACHNUS Hille Ris Lambers Lachninae: Lachnini

One species on Elaeagnus in Taiwan, distinguished in particular by very

numerous protuberant secondary rhinaria on the antennae of alatae (Hille
Ris Lambers, 1956c).
880 Sinomegoura

Sinolachnus niitakayamensis (Takahashi) Apterae yellowish-brown, blackish

on dorsal abdomen; BL c. 2.5 mm. On branches of Elaeagnus oldhami in
Taiwan (Takahashi, 1927a). Sexual morphs and life cycle unknown.

SINOMEGOURA Takahashi Aphidinae: Macrosiphini

Six species with head ventrally spinulose and with weakly clavate, calf-shaped
siphunculi and a long CAUDA, living on trees and shrubs in East and South-
east Asia. Accounts are available for Japan (Miyazaki, 1971), China (Tao,
1963), northeast India (Raychaudhuri, 1980) and Java (Noordam, 1986).

Sinomegoura citricola van der Goot Apterae broadly spindle-shaped, shiny

dark brown to reddish-brown with bright red eyes, siphunculi pale at bases
and a long dark CAUDA; BL 1.8-2.7 mm. On undersides of leaves or young
growth of numerous tropical shrubs and some trees, in about 20 families.
Distributed throughout East and Southeast Asia, and also in Australia
(Queensland). No sexual morphs are known; apparently it is mainly or entirely
anholocyclic. Various karyotypes have been reported; 2n = 12 (Kulkami,
1984), 2n = 16 (Chen and Zhang, 1985b) and 2n = 18 (Kar and Khuda-
Bukhsh, 1986).

Sinomegoura nepalensis Das and Raychaudhuri Apterae yellowish, with

siphunculi basally pale and distally dark as in citricola, but with a pale
CAUDA; BL 2.8-2.9 mm. On young shoots of Melia sp. in Nepal (B.C. Das
and Raychaudhuri, 1983). Life cycle unknown.

Sinomegoura photiniae (Takahashi) Apterae green, often with a pinkish

tinge (Japan; Takahashi, 1960c) or 'faded orange' (China; Takahashi, 1936a),
with black siphunculi and green or yellowish CAUDA; BL 2.0-2.8 mm. On
Photinia spp. and Raphiolepis umbellata (Miyazaki, 1971). In China, Japan
and India. Sexual morphs and life cycle unknown. 2n = 18 (Khuda-Bukhsh
and Kar, 1990).

Sinomegoura pyri Ghosh and Raychaudhuri Apterae pale yellowish, with

pale siphunculi and CAUDA; BL c. 2.4 mm. On undersides of young leaves
of Pyrus sp. in Assam, India (A.K. Ghosh and Raychaudhuri, 1968b). Ardisia
sp. is also recorded as a host plant (Raychaudhuri et al., 1980b). Other morphs
and life cycle unknown.

Sinomegoura symplocois (van der Goot) Apterae light reddish-brown, with

black eyes, banded antennae, pale brownish-yellow siphunculi with black
apices, and black CAUDA; BL 2.9-3.0 mm. On undersides of young leaves of
Symplocos sessilifolia and S. javanica in Java (van der Goot, 1917; Noordam,
1986) and Malaya (BMNH colln, leg. R. Takahashi). Life cycle unknown.
 Sitobion 881

SINONIPPONAPHIS Tao Hormaphidinae: Nipponaphidini

Two species in East Asia in which the dorsum of the aleyrodiform aptera is
densely covered with hair-like processes.

Sinonipponaphis formosana (Takahashi) Apterae blackish-brown, with a

small yellowish-brown patch on 'margin near caudal apex' (Takahashi, 1927b);
BL c. 1.7mm. Alata black with brownish abdomen; secondary rhinaria
distributed ANT III 32-36, IV 18-20, V 10-13. Apterae are fixed firmly to
branches of Machilus sp. (Takahashi, 1929) in Taiwan and China (Szechuan;
Tao, 1966). Life cycle unknown.

Sinonipponaphis hispida Noordam Apterae dull black or brownish-black

with reddish-brown CAUDA, without wax; BL 0.8-1.0 mm. Living scattered
on undersides of leaves of Lithocarpus indutus in Java (Noordam, 1991).
Alatae and life cycle unknown.

SIPHONATROPHIA Swain Aphidinae: Aphidini

One species which is a specialized feeder on Cupressaceae in USA and Central

America (Remaudière and Muñoz-Viveros, 1991).

Siphonatrophia cupressi (Swain) Apterae pale green (concolorous with

leaves of host), body strongly domed dorsally and flattened against leaf
ventrally; BL 1.0-1.7 mm. Alatae have forewing media 1- or 2-branched. On
Cupressaceae (Juniperus, Cupressus, Widdringtonia) in western USA, Mexico
and Central America (Honduras). Monoecious holocyclic; oviparae and both
alate and apterous males in October-November (Palmer, 1952).

SITOBION Mordvilko Aphidinae: Macrosiphini

About 85 species resembling Macrosiphum but with less well-developed

antennal tubercles and often with a more sclerotic dorsum and shorter hairs.
The genus is hard to define; the karyotype (2n = 16 or 2n = 18) suggests that
the strong similarity to Macrosiphum (which have 2n = 10) may be due to
morphological convergence. There is a broad range of host plant relationships,
882 Sitobion

with about 20 species recorded partly or entirely from trees. Sitobion are
distributed throughout the northern hemisphere and have also undergone an
adaptive radiation in Africa south of the Sahara. See also B & E, 1984,
pp.349-350.

Sitobion africanum (Hille Ris Lambers) Apterae variably coloured, yellow-

ish-green to reddish or greenish-brown, the reddish or brownish forms usually
being densely powdered with wax; BL 1.6-3.1 mm. Mainly on Gramineae (see
B & E, 1984, p. 350), but also on numerous dicots including some trees (e.g.
Dombeya, Ficus, Phyllanthus). Euphorbiaceae-feeders have lateral abdominal
tubercles; possibly more than one species is being identified as africanum. In
Africa, southwest Asia (Syria) and possibly South America (Chile). Life cycle
unstudied; oviparae were described from Ficus by Müller and Scholl (1958).

Sitobion autriquei Remaudière Apterae shining brown-black, dorsally con-

vex, with black siphunculi and a pale CAUDA with a dusky tip; BL
3.9-4.3 mm. On various dicots, including Laportea aestuans, as yet known
only from Burundi (Remaudière and Autrique, 1985). Life cycle unknown.

Sitobion bambusicola (L.K. Ghosh) Apterae yellowish; BL c. 3.7mm. In

colony under young, tender leaves of an unidentified Bambusa sp. in Himachal
Pradesh, India (L.K. Ghosh, 1986). Other morphs and life cycle unknown.
It apparently differs from S. miscanthi, which occasionally colonizes bam-
boos, by having longer SIPH (2.0-2.1 x CAUDA), compared with 1.4-1.9 x
CAUDA in miscanthi.

Sitobion berchemiae (Takahashi) Apterae deep green with brownish tinge

on head and thorax, black antennae, mainly dark legs, black siphunculi and
a pale yellowish CAUDA; BL 1.3-1.8 mm (Miyazaki, 1980a, as Macrosiphum
berkemiae). On Berchemia lineata in China (Canton; Takahashi, 1938b), and
B. racemosa in Japan. Life cycle unknown. [The Chinese and Japanese
populations are possibly distinct species, as they apparently differ in the
number of accessory hairs on R IV; 8 in berchemiae (Takahashi, 1938b),
4-6 in berkemiae (Miyazaki, 1980a).]

Sitobion burundiense Remaudière Apterae shining dark green with black

antennae and siphunculi, darkish legs and CAUDA; BL 2.1-2.9 mm. It differs
from other African Sitobion in having 4 hairs (not 3) on first tarsal segments.
On Leguminosae, including Cassia sp., in Burundi (Remaudière and Autrique,
1985). Life cycle unknown.

Sitobion congolense Doncaster and Hille Ris Lambers Apterae mid- to dark
green; BL 2.2-2.5 mm. In West Africa, usually on Vernonia spp. (Com-
positae), forming colonies at growing points, but Remaudière and Autrique
(1985) recorded it from Harungana madagascariensis, both in Burundi and
Ivory Coast (cf. S. nigeriense). Parasitism by Aphidius sp. was observed.
Life cycle unknown.
 Sitobion 883

Sitobion cornifoliae (Shinji) Apterae translucent, whitish, with banded

antennae; BL c. 2.2 mm. Alatae pale yellow-green with yellow-brown thorax
and dusky appendages (Moritsu, 1983). On undersides of leaves of Cornus
spp. in Japan and Korea. Presumably monoecious holocyclic; oviparae were
collected in November (Miyazaki, 1971). [An undescribed Macrosiphum sp.
from Cornus stolonifera in British Columbia, Canada may key out to this
species, but is distinct.]

Sitobion halli (Eastop) Apterae uniformly pale green, with black-tipped

siphunculi and pale CAUDA; BL 1.8-2.9 mm. Immatures are somewhat wax-
powdered (Eastop, 1959). On young growth of trees in various families
(Apocynaceae, Rubiaceae, Moraceae, Euphorbiaceae, Melianthaceae, Myrta-
ceae) in Central and southern Africa (Burundi, Tanzania, Zimbabwe, South
Africa). Life cycle unknown.

Sitobion krahi (Eastop) Apterae green, with siphunculi pale only on basal
third and a pale CAUDA; BL 2.5-3.0 mm. Alatae have siphunculi wholly
black and dark marginal abdominal sclerites, a small antesiphuncular sclerite
and a large postsiphuncular sclerite. Collected on growing points of saplings of
a forest tree, probably Rauwolfia vomitoria but possibly Conopharyngea
chippii, in Ghana (Eastop, 1959). Specimens trapped or on unidentified hosts
have been collected in Nigeria, Sierra Leone and Tanzania (BMNH colln), and
Remaudière and Autrique (1985) recorded krahi from Mangifera indica in
Burundi. Life cycle unknown.

Sitobion leonidasi Remaudière Apterae variably pigmented but usually with

a shiny black abdomen and both siphunculi and CAUDA black; BL 1.8-
2.5 mm. Alatae have dark wing veins and variably developed dark abdominal
markings. On Bersama ugandensis, living particularly in leaf galls formed by
a psyllid. Only known from Burundi (Remaudière and Autrique, 1985).

Sitobion nigeriense (Eastop) Apterae green with dark siphunculi and pale
CAUDA; BL 1.5-2.2 mm. Mostly recorded from Vernonia (Nigeria, Kenya,
Ethiopia; BMNH colln), but Remaudière and Autrique (1985) recorded it
twice colonizing Harungana madagascariensis in Burundi. Parasitism by a
?Trioxys sp. was observed. Life cycle unknown.

Sitobion ochnearum (Eastop) Apterae salmon-pink (but this colour was

possibly due to fungal infection); BL 1.7-2.3 mm (Eastop, 1959). On Ochna
spp. (afzelii, leptoclada, pretoriensis, pulchra) in eastern and southern Africa
(Angola, Kenya, South Africa, Zambia, Zimbabwe). Life cycle unknown.

Sitobion papillatum subnudum Remaudière Apterae matt dark brown, with

dark siphunculi and dusky CAUDA; BL 2.1-2.5 mm. On Arundinaria alpina
in Burundi (Remaudière and Autrique, 1985). Described as a subspecies of
S. papillatum s. str. (which lives on Pennisetum purpureum), but it should
perhaps have full species status. Life cycle unknown.
884 Slaviím

Sitobion phyllanthi (Takahashi) Apterae dirty olive green with dark brown
head and dorsal markings (Müller, 1959; as S. adgnatum); BL 1.5-2.1 mm.
Prepared specimens of apterae have variably developed narrow dorsal abdom-
inal cross-bands between dark marginal sclerites. Described from Phyllanthus
sp. in Mauritius (Takahashi, 1937c). Remaudière (1988) suggested that
5. adgnatum Müller, widely distributed on Euphorbiaceae in Africa, is this
species, and the BMNH collection contains specimens collected by A. van
Harten on Phyllanthus maderaspatensis in Angola. Life cycle unknown.

Sitobion rhamni (Clarke) Apterae yellow-green, with a large round patch of

reddish internal pigment in the centre of the dorsum (Essig, 1917), antennae
ringed with black, pale siphunculi and CAUDA; BL 1.8-2.4 mm. On under-
sides of leaves of Rhamnus spp. in western North America. Life cycle
unknown, except that fundatrices occur in March on Rh. purshiana in British
Columbia (BMNH colln, leg. C.K. Chan). 2n = 16*.

Sitobion salviae (Bartholomew) Apterae and alatae brownish to bottle

green, brick red on head and thorax, with antennal flagellum and siphunculi
black and CAUDA pale (Cermeli, 1970; as Macrosiphum mesosphaeri); BL
1.6-2.3 mm. On woody Labiatae (Hyptis, Inga, Salvia) in Central and South
America (see Smith and Cermeli, 1979). Life cycle unknown.

Sitobion takahashii (Eastop) Colour of apterae in life not recorded in

original description (Takahashi, 1938a; as Macrosiphum phyllanthi), pro-
bably greenish, with black antennae and siphunculi; BL 1.7-2.3 mm. On
Phyllanthus spp. in China, Thailand and India, and perhaps also on Mimosa
pudica in India (M.R. Ghosh et al., 1977a). [Specimens from Phyllanthus
(maderaspatensis, urinaria) from southern India, Java and Singapore in the
BMNH collection have a longer antennal PT and shorter R IV+V than
takahashii s. str., and seem to be the form referred to as a subspecies by
David (1958); possibly the name mimosae Ghosh, Basu and Raychaudhuri
is applicable to this form, but further work is necessary to confirm that
there are indeed two very similar species, both living on Mimosa and
Phyllanthus.]

SLA VUM Mordvilko Pemphiginae: Fordini

Four species closely related to Aploneura, producing sac-like or coral-like galls

on Pistacia spp. in southwest and Central Asia. Three species produce
emigrant alatae and are apparently heteroecious, with a two-year life cycle as
in most other heteroecious Fordini, but their secondary host generations
are still unknown. The fourth is monoecious. Davatchi (1958) revised the
genus.
 Smynthurodes 885

Slavum esfandiarii Davatchi and Remaudière Galls on twigs of Pistacia

mutica in Iran are coral-like, red when mature, with branches slightly larger
than those of S. wertheimae and S. mordvilkoi (Davatchi and Remaudière,
1957). Heteroecious holocyclic; alatae (BL 2.0-2.2 mm) emerge in November
and fly to an unknown secondary host.
Slavum lentiscoides Mordvilko Galls (Fig. 126N, p. 700) are bag-like out-
growths of the leaves of Pistacia spp. (mutica, vera), similar in position to
those of Geoica, but with pointed apices. In southwest and Central Asia (Iran,
Tadzhikistan, Turkmenia). Heteroecious; alatae (BL 1.6-2.2 mm) emerge in
October and migrate to an unknown host. It is possible that S. lentiscoides
is the primary host form of Aploneura ampelina, which lives on roots of vines
and grasses in southern Europe, Africa and southwest and Central Asia, but
experimental work is needed to confirm this.
Slavum mordvilkoi Kreutzberg Galls on twigs of Pistacia vera are coral-like,
similar to those of S. wertheimae. Recorded only from wild P. vera, in
Turkmenia, Afghanistan and eastern Iran (Kreutzberg, 1953). Apparently
heteroecious, but only the primary host part of the life cycle is known.
Slavum wertheimae Hille Ris Lambers Galls are branching, coral- or
cauliflower-like (Fig. 126O, p. 700), reddish when mature, 5-15 cm in dia-
meter, on twigs and branches of Pistacia atlantica and P. mutica in Iran, Israel
and Syria. Monoecious holocyclic, presumably with a one-year cycle; alatae
(BL 1.7-2.2 mm) emerging from galls in September are sexuparae and pre-
sumably give birth to sexuales on the trunk, although this has not been
recorded. Aloni et al. (1989) studied the differentiation of plant tissue in the
galls and Wool and Manheim (1983) studied morphological variation within
the galls.

SMYNTHURODES Westwood Pemphiginae: Fordini

One species characterized by the elongate second antennal segment and the
thick sclerotic rims of the primary sensoria on the last two segments.
Smynthurodes betae Westwood Galls on Pistacia spp. (atlantica, mutica
and, rarely, vera) are yellow-green or red, spindle-shaped, about 20 mm long,
formed by rolling of the edge of the leaflet near its base (Fig. 126P, p. 700).
These are secondary galls, produced by the progeny of the fundatrix, which
lives in a small red mid-rib gall (Burstein and Wool, 1991). Heteroecious
holocyclic, with a two-year cycle; alatae (BL 1.3-1.6 mm) emerge in September-
November and migrate to roots of numerous, mostly dicotyledonous, plants.
The holocycle is recorded throughout the range of the primary hosts; Algeria,
Morocco, Israel, Syria, Iran, southern Crimea and Transcaucasus (Mordvilko,
886 Sorbaphis

1935, as Trifidaphis phaseoli; Davatchi, 1958). Anholocyclic populations

occur commonly on secondary hosts throughout the world (see B & E, 1984,
p. 356). Wool and Burstein (1991a, b) studied phenology and ecology of gall
generations and interactions with a parasitoid in Israel. 2n = 8.

SORBAPHIS Shaposhnikov Aphidinae: Macrosiphini

One species related to Sappaphis, but with 4 instead of 3 hairs on 1st segments
of fore and mid-tarsi.

Sorbaphis chaetosiphon Shaposhnikov Spring colonies curl leaves of Sorbus

spp. downward and turn them reddish. Heteroecious holocyclic; emigrant
alatae (BL 2.4-2.8 mm, with shining black abdomen due to an extensive
sclerotic patch), migrate in June to Ligularia or Polygonum (Pashchenko,
1988b), where they live in ant shelters on stems or at bases of leaves. In Japan,
Korea, China, Siberia (lower course of the Amur and maritime region) and
the southern Urals. 2n = 38.

STEGOPHYLLA Oestlund Drepanosiphinae: Phyllaphidini

About six species on nearctic Quercus. Alate females apparently do not occur
and males may be apterous or alate within a species. Hille Ris Lambers (1966a)
keyed the species then known and discussed the confused nomenclature.

Stegophylla essigi Hille Ris Lambers Apterae are broadly oval, pale grey-
green to olive, covered in white wax wool; BL 1.2-1.8 mm. Feeding by
fundatrices in spring causes edges of young leaves to curl upward and
become thickened and red. Monoecious holocyclic on native Quercus spp.
in California, USA, with oviparae and both apterous and alate males in
November, but populations may continue parthenogenetically on evergreen
oaks such as Q. agrifolia, especially in leaves spun together by caterpillars
(Hille Ris Lambers, 1966d). 2n = 12.

Stegophylla mugnozae Remaudière and Quednau Apterae are broadly pear-

shaped, creamy white, covered with white or bluish-white wax wool; BL
1.4-1.8 mm. On leaves of Quercus crassipes and Q. castanea in Mexico
(Remaudière and Quednau, 1985). Other morphs and life cycle unknown.

Stegophylla querci (Fitch) (= quercicola Baker nec Monell) Apterae are

oval, pale greenish- or brownish-yellow, covered in dense white wax wool; BL
 Stomaphis 887

1.0-1.5 mm. In small, scattered colonies on upper and undersides of leaves of

Quercus spp., widely distributed in North America (see Smith and Parron,
1978). Monoecious holocyclic; oviparae and both apterous and alate males
occur in September-October (Hottes and Frison, 1931; as quercicola Baker).

Stegophylla quercicola (Monell) (= quercina Quednau) Apterae pale

yellowish-green with brown head, covered with bluish-white wax wool; BL
1.6-1.9 mm. On leaves of Quercus spp., in dense flocculent masses which
may cause curling and yellowing (Quednau, 1966). Monoecious holocyclic;
oviparae and both apterous and alate males in October. In eastern North
America. 2n > 30*.

Stegophylla quercifoliae (Gillette) Apterae yellowish to yellowish-green with

brown head, wholly covered in white wax wool; BL 1.1-1.5 mm. On leaves
of Quercus spp., especially Q. gambellii, large colonies causing leaf curl.
Monoecious holocyclic, with oviparae and alate males in October. In western
USA (California, Colorado, Utah, Idaho).

STOMAPHIS Walker Lachninae: Lachnini

About 25 palaearctic species of very large, stem- or root-feeding aphids,

remarkable for their very long rostrum which enables them to probe down
through crevices in thick bark of oak and other trees. All are probably
monoecious holocyclic; males are small, apterous and lack mouthparts.
Colonies are invariably attended by ants (Lasius). Protaphidius wissmannii
is a specialized parasitoid. Accounts and keys are available for Europe
(Heinze, 1962; Szelegiewicz, 1962; Czylok and Blackman, 1991), Ukraine
(Mamontova, 1972) and Japan (Takahashi, 1960b; Sorin, 1979a).

Stomaphis aceris Takahashi Colour in life not recorded; BL of aptera

c. 4.5 mm. On Acer sp. in Japan (Takahashi, 1960b). Biology and sexual
morphs unknown.

Stomaphis alni Sorin Apterae dark brown; BL c. 6mm. On Alnus hirsuta

in Japan. Sexual morphs in September (Sorin, 1965).

Stomaphis aphananthae Sorin Apterae blackish-brown; BL c. 4.7 mm. On

Aphananthe aspera in Japan. Sexual morphs in November (Sorin, 1979a).

Stomaphis asiphon Szelegiewicz Colour in life unknown; BL of aptera

4.8-5.9 mm. Described from roots of an undetermined tree species in Korea
(Szelegiewicz, 1975). Pashchenko (1988b) recorded its host in eastern Siberia
as Juglans mandshurica. Sexual morphs undescribed.
888 Stomaphis

Stomaphis betulae Mamontova Apterae shining blackish-brown; BL

c. 5.3mm. In bark crevices on Betula spp. in Ukraine (Mamontova, 1972).
Sexual morphs undescribed.

Stomaphis bratislavensis Czylok and Blackman Apterae brownish, coated

with powdery wax; BL 4.3-4.8 mm. On bark of Quercus petraea in former
Czechoslovakia. Sexual morphs in October (Czylok and Blackman, 1991).
2n = 8 (Blackman, 1990).

Stomaphis carpini Sorin Colour in life unrecorded; BL of aptera c. 5.5 mm.

On Carpinus tschonoskii in Japan (Sorin, 1965). Sexual morphs undescribed.

Stomaphis cupressi Pintera Apterae yellowish-brown, possibly wax-dusted;

BL 5.0-5.7 mm. Described from specimens collected on trunk of Cupressus
benthami in Kenya (Pintera, 1965), and since found on C. macrocarpa
in France (BMNH colln, leg. G. Remaudière) and on Cupressus sp. in
Italy (BMNH colln, leg. D. Roberti). Sexual morphs unknown. 2n = 14.
[Mamontova (1982) described a population on C. sempervirens in Abkhazia
(Caucasus) as a subspecies, 5. cupressi caucasica.]

Stomaphis fagi Takahashi Colour in life not recorded; BL of aptera

c. 5.3mm. On Fagus sp. in Japan (Takahashi, 1960b). Biology and sexual
morphs unknown.

Stomaphis graffii Cholodkovsky Apterae covered in white wax powder; BL

5-6mm. On trunks of Acer spp., usually campestre, often at or below
soil level, attended by Lasius brunneus. In Europe (France, Poland, former
Czechoslovakia, Rumania). Sexual morphs occur in October. A population in
Ukraine was described by Mamontova-Solukha (1963, 1972) as a subspecies,
S. graffii acerina. [Tashev (1961) noted a species resembling S. graffii on
Populus nigra in Bulgaria.]

Stomaphis japonica Takahashi, Plate 15f Apterae shiny brown, sometimes

wax-dusted; BL c. 5 mm. On trunk of Quercus spp. (acutissima, mongolica)
in Japan (Takahashi, 1960b; as 5. quercus japonica). Sexual morphs in
October-November. Kamiju and Takada (1983) studied a parasitoid and
hyperparasitoid of this species. 2n = 10 (Blackman, 1990).

Stomaphis liquidambaris Takahashi Apterae blackish; BL c. 5.2mm. On

stems of Liquidambar formosana in Taiwan (Takahashi, 1925). Sexual
morphs unknown.

Stomaphis longirostris (Fabricius) Apterae powdered with greyish-white

wax; BL 6.0-6.5 mm. On trunks of Populus spp. (nigra, tremula) and Salix
spp. (alba, viminalis). In continental Europe (France, Germany, Italy, The
Netherlands, Spain) and western Siberia. Tashev's (1961) record from Quercus
sp. in Bulgaria may be an error; however, the species resembling S. graffii
 Stomaphis 889

that he found on P. nigra is possibly longirostris. We have no record of the

sexual morphs.

Stomaphis malloti Sorin Apterae dark brown; BL c. 5.2mm. On Mallotus

japonicus in Japan. Sexual morphs in October-November (Sorin, 1979a).

Stomaphis mordvilkoi Hille Ris Lambers Apterae dark brown; BL 5.3-

7.3mm. On trunk of Juglans regia; described from West Bengal, India
(Hille Ris Lambers, 1933b) and subsequently rediscovered in northern Italy
(Colombo, 1982). Sexual morphs unknown.

Stomaphis pini Takahashi Colour in life unknown, probably blackish; BL

c. 4.5mm. On basal part of trunk or roots of Pinus densiflora in Japan
(Takahashi, 1960b; Inouye, 1970). Sexual morphs not recorded.

Stomaphis quercus (Linnaeus), Plate 15e Apterae elongate oval, shining

dark brown; BL 5.5-7.0 mm. In bark crevices on trunk of Quercus spp.
(robur, petraea); also on Betula pendula and occasionally Alnus glutinosa.
Throughout Europe. Sexual morphs in October-November. Goidanich (1958)
made a detailed study of its life cycle and relation with the ant Lasius
fuliginosus. 2n = 10 (one sample with 2n = 8, suggesting that more than
one species may be involved; see Blackman, 1990).

Stomaphis radicicola Hille Ris Lambers Apterae broadly oval, powdered

with grey dust; BL 4.5-6.0mm. On roots of Betula spp., deep underground.
Only known from The Netherlands (Hille Ris Lambers, 1947b). Sexual
morphs unknown.

Stomaphis rhusivermiciflua Zhang Appearance in life unknown; BL "of

aptera c. 5.4mm. On Rhus verniciflua ( = vernicifera) in Yunnan, China
(Zhang and Zhong, 1985e). Other morphs and biology unknown.

Stomaphis sinisalicis Zhang Aptera without wax powder; BL c. 6.0 mm. On

Salix matsudana, collected at Beijing, China (Zhang and Zhong, 1982b).
Other morphs and biology unknown.

Stomaphis takahashii Sorin Appearance in life not recorded; BL of aptera

c. 5.0mm. On Acer pictum var. typicum and Betula latifolia in Japan
(Hokkaido). Sexual morphs in September (Sorin, 1965).

Stomaphis ulmicola Inouye Apterae shiny brown-black; BL c. 6.3 mm. On

trunk of Ulmus propinqua ( = japonica) in Hokkaido, Japan (Inouye, 1938;
Takahashi, 1960b). Other morphs unknown.

Stomaphis yanonis Takahashi Apterae grey-brown with 2 rows of dark

brown dorsal patches; BL c. 5.0mm. In crevices on bark of trunk of Celtis
sinensis and Zelkova serrata, in Japan, Korea, China and Taiwan. Takahashi
890 Subtakecallis

(1960b) described a population on Aesculus turbinata as a subspecies,

S. yanonis aesculi. Sexual morphs in October-November (Sorin, 1965).
2n = 20 (Honda, 1921) or 16? (Blackman, 1990).

SUBTAKECALLIS Raychaudhuri and Pal

Drepanosiphinae: Phyllaphidini

Two bamboo-feeding species in India, resembling Takecallis in the anterior

projection of the clypeus, but differing in that the ANT PT/BASE ratio is
only 0.24-0.55 and the tibiae are evidently glandular.

Subtakecallis brevisetosus Raychaudhuri and Pal All viviparae alate, appear-

ance in life not recorded, probably wax-covered; BL 1.9-2.3 mm. On uniden-
tified bamboo in West Bengal, India (Raychaudhuri and Pal, 1974). Life cycle
unknown.

Subtakecallis pilosa (David, Rajasingh and Narayanan) All viviparae

alate, covered in white powdery wax, with brown antennae, legs and dorsal
abdominal sclerites; BL 2.3-2.6 mm. On Bambusa sp(p). in northeast India
(Meghalaya, West Bengal). Oviparae and males were described by Agarwala
and Mahapatra (1990).

SUMATRAPHIS Takahashi Greenideinae: Cervaphidini

One oriental species on Ulmaceae with flattened apterae having corrugated

dorsal cuticle and protruding spiracles.

Sumatraphis celti Takahashi Apterae dull green (in Sulawesi; J.H. Martin,
pers. comm.) or yellow to deep brown (in India; A.K. Ghosh et al., 1971e);
BL 1.5-1.7 mm. On new growth of Celtis tetrandra or ?Trema orientalis;
the apterae are flattened into angles of new shoots (J.H. Martin, pers.
comm.). In Indonesia (Sulawesi, Sumatra) and India (West Bengal). Alatae
of Sumatraphis have also been trapped in Nepal (BMNH colln, leg. K.C.
Sharma). The alate morph has not been properly described, and sexual morphs
and life cycle are unknown. The colour difference noted between Indonesian
and Indian specimens suggests that two species might be involved; the host
in India was unidentified.
 Symydobius 891

SYMYDOBIUS Mordvilko Drepanosiphinae: Phyllaphidini

Six species of medium to large, rather shiny dark brown aphids associated with
Betula and Alnus, forming colonies on 'the branches and twigs, invariably
attended by ants. A seventh species will shortly be described from Betula nana
in the Czech Republic (J. Holman, pers. comm.). All the apterae (even the
oviparae) are 'alatiform', with a similar pattern of sclerotization and pigmenta-
tion to that of the fully-winged adults. Monoecious holocyclic; males are
apterous, and oviparae have the posterior abdominal segments extended into
an ovipositor-like structure. Trioxys betulae is a specialized parasitoid.

Symydobius alniarius Matsumura Reddish or yellowish-brown, with dark

brown cross-bands and often with white powder on posterior segments; BL
3.1-3.4 mm. On stems and twigs of Alnus hirsuta, A. hirsuta var. sibirica and
A. japonica in Japan (Higuchi, 1972), Korea and eastern Siberia (Primorskiya
Kray). Sexual morphs have not been described. 2n = 20.

Symydobius americanus Baker Shiny dark brown to black; BL 2.4-2.7 mm.

On stems and twigs of Betula papyrifera in northern USA and Canada. Sexual
morphs in September-October (Baker, 1918).

Symydobius intermedius Gillette and Palmer Olive green to dark grey-

brown; BL 2.2-2.6 mm. On stems and twigs of Betula occidentalis in western
USA (Colorado, Utah, Idaho) and also recorded from British Columbia.
Sexual morphs in September-October (Gillette and Palmer, 1931). 2n = 16.

Symydobius kabae Matsumura Dark brown; BL 2.5-3.0 mm. On bark of

last year's branches and twigs of Betula maximowicziana and B. platyphylla
var. japonica in Japan (Higuchi, 1972); also recorded from B. gmelini
in Mongolia (as alniarius - see Holman and Szelegiewicz, 1974a), from B.
platyphylla var. latifolia and Alnus (hirsuta var. sibirica, japonica) in Korea
and from B. dahurica in eastern Siberia (Primorskiya Kray). Paik (1972)
recorded apterous oviparae in Korea in November. 2n = 26? (Shinji, 1931;
but see Blackman, 1986).

Symydobius minutus Quednau and Shaposhnikov Yellowish-brown, with

dark transverse bars and white wax powder posteriorly; BL 2.2-2.4 mm.
On shoots of Alnus japonica in eastern Siberia (Primorskiya Kray) and Korea
(Quednau and Shaposhnikov, 1988). Sexual morphs unknown.

Symydobius oblongus (von Heyden) Shiny dark brown; BL 2.0-3.5 mm. On

twigs, young stems and branches of Betula spp. (nana, pendula, pubescens)
throughout Europe and across Asia; the form found in eastern Siberia
892 Taiwanaphis

(Quednau and Shaposhnikov, 1988) is possibly the subspecies alni Bozhko

(1979). Sexual morphs in October-November (Heie, 1982). Holman (1990)
studied developmental morphology of fundatrices and their progeny, and
Fowler and Macgarvin (1985) studied the effects of ant attendance on popula-
tion growth. See Heie (1982) and Stroyan (1977). 2n (female) = 15, 2n
(male) = 14 (Blackman, 1988).

TAIWANAPHIS Takahashi Drepanosiphinae: Phyllaphidini

Twelve species on trees in various families, but especially Myrtaceae, in East

and Southeast Asia, or on Nothofagus in Australasia and New Zealand. In
several species only alate viviparae are known, but this may be because
apterous viviparae are well camouflaged on the stems and easily overlooked,
whereas the alatae develop on the leaves. Accounts are available for Java
(Noordam and Hille Ris Lambers, 1985) and India (A.K. Ghosh and Quednau,
1990). Carver (1978) keyed the species (subgenus Sensoriaphis) on Nothofagus.

Taiwanaphis atrovirens Noordam and Hille Ris Lambers Alatae have head,
pronotum and sides of abdominal segments 3-8 yellowish or brownish,
mesonotum and central part of dorsal abdomen dark green; BL 1.2-1.5 mm.
Immature alatae are yellowish-white with a dorsal greenish-black area.
Apterous viviparae, other morphs and life cycle unknown. On upper and
lower sides of new leaves and developing shoots of ?Syzygium sp. in Java
(Noordam and Hille Ris Lambers, 1985).

Taiwanaphis atuberculata Noordam and Hille Ris Lambers Alatae are

orange with grey sides to thorax and abdominal segments 1-5, banded anten-
nae and black-bordered forewing veins; BL 1.5-1.7 mm. Other morphs and
life cycle unknown. On upper sides and undersides of leaves and sometimes
on developing shoots of Syzygium lineatum in Java.

Taiwanaphis decaspermi Takahashi Apterae and alatae brownish-black to

black; BL 1.2-1.6 mm. Alatae have thickly bordered forewing veins. Described
from Decaspermum fruticosum in Taiwan, attacking undersides of young
leaves and shoots. Subsequently recorded from other Myrtaceae (Eugenia
minutiflora, Syzygium buxifolium) in China (Kwantung, Hong Kong). Alate
males were collected in Hong Kong in March (BMNH colln, leg. Com-
monwealth Institute of Entomology).

Taiwanaphis (Sensoriaphis) furcifera Carver and White Alatae have pale

green abdomen, head and pronotum with two longitudinal dark stripes, and
mainly black antennae and legs; BL 1.8-2.8 mm. On tender young shoots
and suckers of Nothofagus moorei in New South Wales, Australia. Carver and
 Taiwanaphis 893

Hales (1974) described all morphs and gave life cycle details; monoecious
holocyclic, with oviparae and alate males occurring concurrently with alate
viviparae through spring, summer and autumn. Eggs are flat, colourless and
wax-covered, and are laid on leaves.

Taiwanaphis kalipadi (Raychaudhuri and Ghosh) Colour in life not recorded;

aptera rather elongate oval, BL 1.2-1.5 mm. On Annona squamosa in West
Bengal, India (Raychaudhuri and Ghosh, 1964). Other morphs and biology
unknown, but possibly T. randiae, known only from alatae, is this species.

Taiwanaphis memecyloni Noordam and Hille Ris Lambers Alatae are

yellow with black sides to thorax and grey-bordered forewing veins; BL
1.0-1.5 mm. On upper surfaces of young leaves, with immatures positioned
close against the main vein, on Memecylon spp. in Java (Noordam and Hille
Ris Lambers, 1985). Other morphs and life cycle unknown.

Taiwanaphis montanicola Noordam and Hille Ris Lambers Apterae and

alatae brown, the alatae having black-bordered forewing veins; BL 1.7-
2.2mm. On undersides of young leaves of Syzygium racemosum in Java
(Noordam and Hille Ris Lambers, 1985). Life cycle unknown.

Taiwanaphis (Sensoriaphis) niuginii Carver Both apterae and alatae occur,

colour in life not recorded; BL of aptera 1.5-2.2 mm, of alata 2.0-2.4 mm.
On undersides of young leaves of Nothofagus carrii in Papua New Guinea
(Carver, 1978). Life cycle unknown.

Taiwanaphis (Sensoriaphis) nothofagi Cottier Apterae yellowish-green to

brownish; BL 1.1-1.5 mm. Well camouflaged, feeding on young stems of
Nothofagus spp. in New Zealand. Alatae become mature on the undersides
of leaves. Monoecious holocyclic, with oviparae and alate males in September
to February (BMNH colln, leg. VFE). 2n = 10.

Taiwanaphis pseudocaudata Noordam and Hille Ris Lambers Alatae have

head and prothorax yellow, mesothorax and abdomen orange with grey sides;
BL 1.1-1.6 mm. Along veins on upper and lower sides of young leaves and
shoots of Syzygium syzygioides in Java (Noordam and Hille Ris Lambers,
1985). Other morphs and life cycle unknown.

Taiwanaphis randiae Ghosh, Banerjee and Raychaudhuri Alatae yellowish

with dark markings and heavily bordered forewing veins; BL 1.7-2.0 mm. On
upper and lower sides of terminal leaves of Randia sp. in India (Arunachal
Pradesh), attended by ants (A.K. Ghosh et al., 197Id). Other morphs and life
cycle unknown; possibly a synonym of T. kalipadi, which is only known from
apterae.

Taiwanaphis (Sensoriaphis) tasmaniae Carver and Martyn Apterae have

dirty yellow head and prothorax, rest of body olive green and rather shiny;
894 Takecallis

BL 1.5-2.0 mm. On stems of young shoots of Nothofagus cunninghami in

Tasmania (Carver and Martyn, 1962). The alatae morph was described by
Carver and Martyn (1965). Life cycle unknown.

Taiwanaphis sp. An as yet undescribed species of Taiwanaphis was collected

on Melaleuca spp. (lanceolata, teretifolia) in Western Australia (Carver, 1978,
footnote; BMNH colln, leg. J.C. Taylor).

TAKECALLIS Matsumura Drepanosiphinae: Phyllaphidini

Five species of narrow-bodied aphids on bamboos, resembling Myzocallis

but with the clypeus bearing an anteriorly directed tubercle, and with ANT
PT/BASE about 1. Usually all viviparae are alate. The genus is oriental in
origin, but several species are now widely distributed. Higuchi (1968) revised
the species then known and accounts are available for Japan (Higuchi, 1972),
India (A.K. Ghosh and Quednau, 1990) and for species introduced into Britain
(Stroyan, 1977). See also B & E, 1984, p. 357.

Takecallis affinis L.K. Ghosh Alatae blackish, wax-covered in life; BL

2.1-2.5 mm. On undersides of leaves of Bambusa sp. and Arundinaria
jounsarensis in Himachal Pradesh, India. Apterous oviparae and alate males
occur in October (L.K. Ghosh, 1986). Takecallis himalayensis, described from
unidentified bamboos in Uttar Pradesh (Chakrabarti, 1988), is clearly a
synonym, to judge from its redescription by A.K. Ghosh and Quednau (1990),
which includes oviparae with the same collection data as paratypes of affinis.
B.C. Das and Chakrabarti (1989b) recorded a new parasitoid species (Trioxys
takecallis from T. himalayensis (= T. affinis) in Garwhal, western Himalaya.

Takecallis arundicolens (Clarke) Alatae pale yellow or greyish-yellow, with

dark dorsal abdominal markings and a black CAUDA; BL 1.8-2.8 mm. [An
unpigmented population with a pale CAUDA occurred at Wisley, UK, in 1967
(BMNH colln, leg. K.M. Harris).] On mature leaves of bamboos (Arun-
dinaria, Bambusa, Phyllostachys, Sasa) in China, Japan, Korea, eastern
Russia and introduced into Europe and USA (California). Life cycle in
East Asia unknown; anholocyclic where introduced, although Leclant (1966)
reported oviparae in southern France. 2n = 18.

Takecallis arundinariae (Essig) Alatae whitish, pale yellow or greyish-

yellow, with paired black elongate spots on ABD TERG 1-7 and a pale
CAUDA; BL 1.7-2.4 mm. An apterous morph has been described, but only
from high altitudes in Taiwan. On undersides of mature leaves of bamboos
(especially Arundinaria and Phyllostachys, but sometimes also Bambusa and
Dendrocalamus). In India, China, Taiwan, Korea, Japan and introduced to
 Taoia 895

England, New Zealand and North America. Life cycle unknown in East Asia;
anholocyclic where introduced. Coffelt and Schultz (1990) studied its popula-
tion dynamics in USA. 2n = 18.

Takecallis himalayensis Chakrabarti See T. affinis.

Takecallis sasae (Matsumura) Alatae green; BL c. 1.8mm. On leaves of

Phyllostachys, Pleiobtastus, Sasa and possibly also Bambusa, in Japan
(Higuchi, 1968). Tao (1964) had T. taiwanus in China and Taiwan according
to Higuchi (1968). Life cycle unknown.

Takecallis taiwanus (Takahashi) Alatae pale green with a pale CAUDA; BL

1.4-2.3 mm. In young, still unrolled leaves and on new shoots of bamboos
(Arundinaria and Phyllostachys, sometimes Bambusa, Sasa). In China,
Taiwan, Japan and introduced to Europe, South Africa, New Zealand and
USA. Cottier's (1953) account under T. arundinariae should be referred to this
species. Life cycle and sexual morphs unknown; probably anholocyclic where
introduced. 2n = 16.

TAOIA Quednau Drepanosiphinae: Phyllaphidini

Two oriental species on Betulaceae, possibly related to Euceraphis and

Cepegillettea, characterized by the small rounded secondary rhinaria, ANT PT
a little shorter than BASE VI and siphunculi with swollen bases and a single
appended hair. Both apterous and alate viviparae occur. Quednau (1973) gave
an account of the genus.

Taoia chuansiensis (Tao) Apterae and alatae yellowish-green, the alatae

having darker legs and antennae, and dark-tipped siphunculi; BL 2.9-3.1 mm.
Living solitarily on undersides of leaves of Alnus crematogyne and Betula
spp. in China (Tao, 1964, as Euceraphis chuansiensis). Sexual morphs and life
cycle unknown.

Taoia indica (Ghosh and Raychaudhuri) Apterae pale green; BL 2.7-

4.1mm. Alatae have darker antennae and legs, and dark-tipped siphunculi.
On Alnus nepalensis and Betula alnoides in India and Nepal (A.K. Ghosh
and Quednau, 1990). Oviparae were collected in December and apterous
(or alatoid) males in April, July and December (R.C. Basu and Raychaudhuri,
1980).
896 Tetraneura

TETRANEURA Hartig Pemphiginae: Eriosomatini

About 30 species in which the apterae have 1-segmented tarsi and the alatae
have a simple, unbranched media in the forewing and usually ANT IV much
shorter than ANT V. At least seven species have host alternation between leaf
galls on Ulmus and roots of Gramineae, and the lesser-known species are
described either from Ulmus or grass roots. The elm galls are invariably
stalked, pouch-like outgrowths on upper sides of leaves. Tetraneura seems to
be predominantly oriental and eastern palaearctic but T. ulmi appears to be
native to northwest Europe, T. caerulescens and T. africana are around the
Mediterranean and in southwest Asia. T. nigriabdominalis is probably east
Asian although holocyclic populations of this species are widespread. Hille Ris
Lambers (1970b) revised the world fauna and there are more recent accounts
from India (A.K. Ghosh, 1984b), China (Zhang et al., 1991) and Japan
(Akimoto, in prep.), but the genus is still in need of further revision. The
karyotype variation may be useful in this respect. (See also B & E, 1984,
pp. 358-361.)

Tetraneura aequiunguis Zhang and Zhang Only the fundatrix is described,

from an unidentified Ulmus sp. in China (Zhang et al., 1991).

Tetraneura africana van der Goot Galls on Ulmus have not been identified,
but Mordvilko (1935) recorded sexuparae collected on trunks of U. campestris
( = procera) in southern Italy.

Tetraneura asymmachia Zhang and Zhang Described from alatae (BL

c. 2mm) obtained from galls on Ulmus pumila and Ulmus sp. in June-July
in China (Zhang et al., 1991). Appears closely related to T. chinensis.

Tetraneura caerulescens Passerini Galls are stalked, hazelnut- or walnut-

sized (Fig. 134G), red when mature, on Ulmus spp. (procera, montana) in
southern and southeastern Europe, North Africa and southwest Asia. Alatae
(BL 1.5-2.2 mm) emerge in late May-July and migrate to found colonies on
roots of Gramineae (Poa annua, Cynodon dactylon). Records of secondary
host populations in China (Zhang et al., 1985) need further confirmation.

Tetraneura chinensis Mordvilko Gall is apparently rather large, sac-like,

broader than high, green and slightly hairy (Fig. 134F). Recorded from Ulmus
sp. in China (Mordvilko, 1935, as T. hirsuta) and U. pumila in Mongolia
(Holman and Szelegiewicz, 1974a). Alatae (BL 2.2-2.6 mm) were obtained
from galls in May-June. Life cycle is unknown.

Tetraneura fusiformis Matsumura Galls on leaves of Ulmus japonica in

Japan are upright spindle-shaped, up to 12mm in height, petiolate at base,
 Tetraneura 897

with numerous granular protuberances and short white silky hairs, usually
crimson when mature. Alatae (BL 1.7-2.3 mm) emerge in June-July through
a lateral opening. Hille Ris Lambers (1970b) redescribed the gall generations
(as nigriabdominalis). The life cycle has not been fully investigated. 2n = 18
(S. Akimoto, pers. comm.).

Tetraneura nigriabdominalis (Sasaki) = akinire Sasaki (Akimoto, in prep.),

Plate 5f Galls are stalked, hairy, elongate, spindle- or pouch-shaped, 15-40
mm high, usually with a pointed apex (Fig. 134E), usually bicoloured green
and rose-red when mature. On various Ulmus spp. (japonica, parvifolia,
procera) in southern and southeastern Europe, Japan, Korea and also in
eastern USA on U. carpinifolia (Stoetzel, 1987; as akinire). Alatae (BL
1.4-2.3 mm) leave the gall through lateral slits in May-July to found colonies
on roots of Gramineae (Tanaka, 1961). Enoki (1986) studied resistance to this
aphid in U. japonica. Hille Ris Lambers' (1970b) descriptions of gall genera-
tions under this name (his 'Group I material') should be referred to T. fusi-
formis (S. Akimoto, in prep.). 2n = 18 (in gall generations; anholocyclic
populations are recorded with '2n'=17, and 19, D. Hales, pers. comm.).

Tetraneura paiki Hille Ris Lambers Described from a single alate sexupara
trapped in Korea (Hille Ris Lambers, 1970b). Moritsu's (1983) account and
illustrations of an aphid under this name in Japan should be referred to
T. nigriabdominalis (Akimoto, in prep.).

Tetraneura polychaeta Hille Ris Lambers Galls are conical, bag-like

(Chakrabarti et al., 1985), on Ulmus laevigata (= villosa) in Pakistan. Alatae
(BL 2.0-3.2 mm) were collected from galls in April, but probably emerge later
in spring. A sexuparae trapped in Korea was also tentatively assigned to this
species (Hille Ris Lambers, 1970b). Life cycle unknown.

Tetraneura radicicola Strand Stalked pouch galls on Ulmus japonica in

Japan are similar to those of T. nigriabdominalis, but hairless or with only
sparse hairs. Alatae (BL 1.7-2.5 mm) migrate in June-July to found colonies
on roots of Gramineae. Presumed anholocyclic populations on grass roots are
recorded from Australia, Malaya, Philippines, Korea, Taiwan, India, Nepal
and Sri Lanka, although on its secondary hosts T. radicicola cannot be
reliably distinguished from T. yezoensis (see A.K. Ghosh, 1984b). 2n = 14.
[Tetraneura persicina, described from alatae (BL c. 2.6mm) from galls on
Ulmus sp. in China (Zhang et al., 1991), appears very similar to radicicola,
but the alatae have fewer dorsal cephalic hairs and their embryos have fewer
spinopleural hairs. See also T. triangula.}

Tetraneura sorini Hille Ris Lambers Galls are stalked, hairy, pouch-
shaped, often somewhat globular, yellow and red (Moritsu, 1983). On Ulmus
spp. (laciniata, japonica, parvifolia) in Japan, Korea and China. Alatae
(BL 1.6-2.0 mm) emerge in June. Secondary host generations unknown.
2n = 16.
898 Tetraneura

Fig. 134. Galls on Ulmus. A, Olegia ulmifoliae on Ulmus japonica (after Pashchenko, 1988b); B, Colopha
compressa on Ulmus sp. (after Mordvilko, 1935); C, C. ulmicola on U. americana (after Hottes and
Frison, 1931); D, Tetraneura ulmi on Ulmus sp. (after Marchal, 1933); E, T. nigriabdominalis on Ulmus
sp. (after Moritsu, 1983, as T. paiki); F, T. chinensis on Ulmus sp. (after Mordvilko, 1935, as
 Thecabius 899

Tetraneura triangula Zhang and Zhang Galls and spring generations

unknown. Described from exules and sexuparae, which are similar to
T. radicicola but much more hairy (W. Zhang and G. Zhang, 1991). Possibly
this species occurs in Japan, on U. japonica (S. Akimoto, pers. comm.).

Tetraneura ulmi (Linnaeus) Galls are stalked, approximately bean-shaped

(Fig. 134D), smooth and shiny, reddish-green and/or yellow, on various
Ulmus spp. in Europe, across Asia to eastern Siberia (Pashchenko, 1988b) and
introduced to North America. Alatae (BL 1.8-2.6 mm) emerge in June-July
to colonize roots of Gramineae. Anholocyclic populations occur commonly
on secondary hosts. Zwölfer (1957) made a detailed study of this aphid.
2n = 14.

Tetraneura yezoensis Matsumura Stalked pouch galls on Ulmus japonica

and U. laciniata in Japan resemble those of T. radicicola. Alatae (BL 1.8-
2.1 mm) migrate in late June-July to colonize roots of Gramineae. Anholo-
cyclic populations on grass roots assigned to this species occur in Australia
(Hille Ris Lambers, 1970b). Akimoto (1990) studied local adaptation to host
plant and possible host-race formation in this species. 2n = 12 in Japan; Chen
and Zhang (1985b) reported 2n = 18 from China, but probably had another
species. [Tetraneura changaica, described from sexuparae only in Mongolia
(Szelegiewicz and Holman, 1980), resembles T. yezoensis but has shorter R
IV+V and different abdominal chaetotaxy.]

THECABIUS Koch Pemphiginae: Pemphigini

About 12 species closely related and very similar morphologically to Pem-

phigus, but with some biological differences, and retained here as a separate
genus despite the reservations of Aoki (1975), with Parathecabius as a sub-
genus. Spring generations form galls by folding the leaves of Populus. The
fundatrix inhabits a small gall set apart from that housing its progeny. Known
secondary hosts are Ranunculus, Lysimachia and Salix. Accounts are available
for North America (Smith, 1974a), northwest Europe (Heie, 1980) and Japan
(Aoki, 1975; as Pemphigus).

Thecabius affinis (Kaltenbach), Plate 6d Galls on Populus spp. in May-

June are formed by folding of young leaves in half along mid-ribs; the outer,

T. hirsuta); G, T. caerulescens on U. procera (after Mordvilko, 1935, as T. rubra); H, Kaltenbachiella

pallida on Ulmus sp. (after Marchal, 1933); I, K. ulmifusa on U. rubra (after Hottes and Frison, 1931,
as Gobaishia); J, K. spinosa on U. japonica (after Akimoto, 1985a); K, K. japonica on U. japonica (after
Akimoto, 1985a); L, Eriosoma grossulariae on U. procera (after Patch, 1913b, as E. ulmi);
M, E. lanuginosum on U. procera (after Marchal, 1933); N, E. flavum on U. carpinifolia (from BMNH specimen).
900 Thecabius

upper side of the leaf becomes blistered and yellowish or reddish. These are
secondary galls containing only the alatiform progeny of the fundatrix, which
inhabits a small gall of its own formed by folding the edge of a leaf (Fig. 131I,
p. 803). Alatae (BL 2.2-3.1 mm, with green abdomen) leave the gall in late
June-July to found waxy colonies at stem bases and on runners of Ranunculus
spp. (Fig. 3). Mordvilko (1935) gave an account of the life cycle and described
all morphs. It occurs on a wide range of Populus spp. throughout Europe and
across Asia to eastern Russia, Japan and Korea. Populations on East Asian
Populus spp. are often treated as a separate species, Th. orientalis Mordvilko,
here regarded as a synonym. Karyotype is 2n = 38 in both England and Japan.
[In North America, Thecabius populiconduplifolius (Cowen) is virtually
indistinguishable from Th. affinis and has an identical biology, but with native
poplars as primary hosts (Palmer, 1952). Harper (1966) and Smith (1974a)
suggested synonymy, but Th. populiconduplifolius is not introduced Th.
affinis, as it has 2n = 28 (Harper and Macdonald, 1966); it is apparently a
good species.]

Thecabius (Parathecabius) gravicornis (Patch) Galls are formed by folding

leaves downward along the mid-rib to form a yellowish-green, swollen pocket
which becomes filled with the developing emigrant alatae. The fundatrix
inhabits a separate small pocket on the upper side of a leaf lamina (Palmer,
1952). On Populus balsamifera in North America, widely distributed but
apparently rather rare; also collected on P. angustifolia in Utah (BMNH
colln, leg. R. Danielsson). Alatae (BL 1.7-2.8 mm, with olive-yellow abdomen)
migrate in July-August, presumably to an unknown secondary host.

Thecabius (Parathecabius) latisensorius Hori Galls like those of Th. gravi-

cornis are found on Populus maximowiczii in Japan and Sakhalin. Alatae (BL
3.0-3.5 mm, with dirty yellow abdomen) migrate in late July-September,
presumably to an unknown secondary host (Hori, 1938; Aoki, 1975). 2n = 18
(Blackman, 1986).

Thecabius (Parathecabius) lysimachiae (Börner) Galls containing develop-

ing emigrant alatae are formed by folding and convoluting the leaf of Populus
nigra (Fig. 131J, p. 803). The fundatrix inhabits a separate oval pocket-gall on
upper side of leaf lamina, as in other Parathecabius. Alatae (BL 2.3-2.6 mm,
with dark, dirty greenish abdomen) migrate in late June-July to roots and
runners of Lysimachia mummularia. Anholocyclic overwintering on Lysima-
chia is common. In Europe and Central Asia; Th. luppovae (Narzikulov),
described from Populus densa in Tadzhikistan, may be a synonym. 2n = 18.

Thecabius populimonilis (Riley) The galls on Populus spp. (angustifolia,

balsamifera, fremontii, trichocarpa) are rows of ovoid, bead-like cells, pale
green or with a reddish tinge, developed from upper side of leaf lamina on
both sides of mid-rib. Each gall in spring contains one developing aptera or
alata. The fundatrices inhabit similar, solitary galls. This species is apparently
facultatively monoecious; Gillette (1913) suggested that the large alatae (BL
 Thelaxes 901

2.7-3.0 mm) produced in individual galls in June-July might recolonize

cottonwoods, but it seems more likely that they found the colonies of apterous
exules in wax on Salix roots observed by Lange (1943) in California. Pre-
sumably alatae produced in these colonies in October are sexuparae and return
to Populus. However, alate sexuparae (BL 1.7-2.0 mm) are also produced in
galls on Populus, 10-12 per gall, in August-October (Maxson and Knowlton,
1929; Palmer, 1952; Harper, 1959a). This interesting life cycle would repay
further study. Widely distributed in USA, western Canada and Mexico.

THELAXES Westwood Thelaxinae

Four oak-feeding species, three in Europe and the Mediterranean region, and
one in North America. Remaudière (1982c) provided a key including charac-
ters of alatae as well as apterae, and Ilharco (1966) keyed first instar nymphs.

Thelaxes californica (Davidson) Appearance in life not recorded, pre-

sumably much like Th. dryophila; BL of aptera 1.2-2.0 mm. On Quercus spp.
in western North America, from British Columbia to Mexico. Monoecious
holocyclic; sexual morphs aestivate in first instar (Davidson, 1917), adult
oviparae occur in September-October (Palmer, 1952). 2n = 12*.

Thelaxes dryophila (Schrank) Apterae broadly oval, rather flattened, dark

brownish-red to purplish-grey with a paler spinal stripe; BL 1.1-2.3 mm.
Alatae have dark marginal sclerites and dorsal cross-bands on posterior
abdominal segments. Ant-attended colonies form at tips of shoots, extending
onto stems, leaf petioles and along mid-ribs on undersides of leaves. On
Quercus spp. in Europe, Mediterranean region and southwest Asia. Mono-
ecious holocyclic; wax-coated sexual morphs aestivate from June to September
on undersides of leaves in forks of veins (Polaszek, 1986). Lombard et al.
(1988) studied composition of the honeydew. 2n = 8.

Thelaxes suberi (del Guercio), Plate 4a, b Apterae vary in colour from pale
green to dark brown; BL 0.9-1.5 mm. Living abundantly on young shoots,
leaves and developing acorns of numerous Quercus spp. (but not robur) in
England, southern Europe, Mediterranean region and southwest Asia. Also
recorded from Castanea sativa (in Italy). Presumably monoecious holocyclic,
but life cycle unstudied; Polaszek (1986) collected presumed sexuparae on
Q. cerris in England in November-December. 2n = 8.

Thelaxes valtadorosi Remaudière Apterae light brown, somewhat waxy;

BL 1.0-1.9 mm. On Quercus spp. in Greece (Remaudière, 1982c). Life cycle
unknown. 2n = 8*.
902 Thoracaphis

THORACAPHIS van der Goot Hormaphidinae: Nipponaphidini

Most species originally described in Thoracaphis have been removed to other

genera, leaving the type species (arboris van der Goot) and a few others,
mostly little known, which can hardly be said to form a homogeneous group.
Probably associated in the past with Distylium as primary host as are other
Nipponaphidini, but now only known from parthenogenetic generations on
what were the original secondary hosts (Quercus, Lindera).

Thoracaphis arboris van der Goot Apterae aleyrodiform, flattened, broadly

oval, bluish-black with wax frosting on sides of body only; BL 1.3-1.6 mm.
Immatures dull greyish-green with rather more wax. On undersides of leaves
of Quercus sp(p). in Java. Alatae collected in May and August (Noordam,
1991). Life cycle unknown; presumably anholocyclic.

Thoracaphis flava Takahashi Apterae aleyrodiform, flattened, broadly

oval, only slightly sclerotized, pale yellowish-brown; BL c. 0.5 mm. On under-
sides of leaves of Quercus sp. in Malaya (Takahashi, 1950). Other morphs
and life cycle unknown; presumed to be anholocyclic. This species clearly does
not really belong in Thoracaphis; possibly it requires a new genus.

Thoracaphis linderae Shinji Apterae aleyrodiform, flattened, broadly oval,

green or brown becoming sooty black with age; BL 1.5-2.0 mm. On twigs of
Lindera spp. in Japan (Shinji, 1941). Alatae were collected in May (Shinji,
1926). Life cycle uncertain; Shinji (1926) reported overwintering as eggs on
Lindera, but this was not confirmed by Takahashi (1958a) in his redescription
of the species.

Thoracaphis sp. on Cinnamomum Apterae aleyrodiform, flattened oval,

reddish-brown with creamy-white markings on prosoma and blackish abdom-
inal plate. On bark of stems of Cinnamomum japonicum in Japan (Moritsu,
1983; as ?cinnamomiae Shinji). This appears to be an undescribed species.

TILIAPHIS Takahashi Drepanosiphinae: Phyllaphidini

Four oriental species very similar to Eucalliplerus, but with larger, pale
siphunculi and more extensive forewing pigmentation. Accounts are available
for Japan (Higuchi, 1972), China (Zhang and Zhong, 1982c) and Siberia
(Pashchenko, 1988b; Quednau and Shaposhnikov, 1988).
 Tiliphagus 903

Tiliaphis coreana Quednau All viviparae alate, greenish-yellow to yellow

with brown-black sides to head and prothorax and longitudinal stripes on
mesothorax, becoming more evident in late summer; BL 2.5-3.5 mm. Fore-
wings have deeply pigmented costal margin and a black zig-zag line between
distal ends of veins. Immatures green with 4 rows of black spots. In rows along
veins on undersides of leaves, or as nymphs on upper sides of leaves, of Tilia
spp. in Korea, China and eastern Siberia. Sexual morphs in September (Paik,
1972, as Tiliaphis japonica; Quednau, 1979; Quednau and Shaposhnikov,
1988). 2n = 38 (Chen and Zhang, 1985b).

Tiliaphis pseudoshinae Quednau All viviparae alate, whitish with brown-

black markings on body and wings as in coreana, but without longitudinal
stripes on mesothorax and with clear base to forewing; BL 2.0-2.6 mm. On
Tilia sp. in Korea and eastern Siberia. Sexual morphs in September (Quednau,
1979; Quednau and Shaposhnikov, 1988).

Tiliaphis shinae (Shinji) All viviparae alate, pale yellow to greenish-yellow

with dark brown markings on body and wings as in pseudoshinae; BL 1.7-
2.4 mm. On undersides of leaves of Tilia spp. in Japan, Korea and eastern
Siberia. Sexual morphs in early October (Takahashi, 1961b; Higuchi, 1972).
2n = 14 (Shinji, 1931).

Tiliaphis shinjii Higuchi All viviparae alate, colour in life not recorded,
dorsal dark markings as in coreana but with paired dark spinopleural patches
on abdominal tergites; BL 2.0-2.7 mm. On Tilia spp. in Japan, Korea and
eastern Siberia. Sexual morphs in September (Higuchi, 1972; Quednau and
Shaposhnikov, 1988).

TILIPHAGUS Smith Pemphiginae: Pemphigini

One North American species, differing from Prociphilus in having alatae

with many more secondary rhinaria on the antennae.

Tiliphagus lycoposugus Smith Large leaf-nest galls are produced in spring

by clumping and cupping terminal leaves so as to resemble a large, poorly-
formed head of lettuce, 10-30 cm in diameter. On Tilia americana in eastern
USA (Maryland, North Carolina). Fundatrices inside galls are brown, globose,
BL 4.1-5.3 mm. Heteroecious holocyclic; emigrant alatae are reddish-brown,
BL 2.6-3.3 mm, leaving gall in June-August to found colonies on roots of
Lycopus virginicus. Return migration of sexuparae to trunks of Tilia occurs
in October-November (Smith, 1965).
904 Tinocallis

TINOCALLIS Matsumura Drepanosiphinae: Phyllaphidini

About 25 mostly small species usually associated with Ulmaceae, although the
three members of subgenus Sarucallis feed on Lythraceae (Lagerstroemia,
Duabanga), and species have also been described from Corylus, Dalbergia
and Sapindus. (Woody Leguminosae seem to be recorded quite regularly as
alternative or 'casual' hosts.) All viviparae are alate and usually have paired
spinal and marginal tubercular processes. Some species have conspicuous
black markings on dorsal body and/or forewings. Problems with the taxo-
nomy of Tinocallis may arise from seasonal variation; alatae in summer and
autumn are often more pigmented than in spring and may have fewer secon-
dary rhinaria. For example, specimens of Tinocallis platani collected in May
have 17-25 (usually 20-23) rhinaria on ANT III, whereas specimens collected
from the same trees in July-October have 7-21 (mostly 11-17) rhinaria on III
(VFE, unpublished data). Richards (1967) keyed the world fauna as then
known and more recent accounts are available for Britain (Stroyan, 1977),
Fennoscandia and Denmark (Heie, 1982), India (Chakrabarti, 1988; A.K.
Ghosh and Quednau, 1990), eastern Russia (Pashchenko, 1988b), China
(Zhang and Zhong, 1980c) and Japan (Higuchi, 1972).

Tinocallis allozelkowae Zhang Appearance in life not recorded (but not

wax-dusted); BL of alata c. 2.7 mm. On Zelkova schneideriana in China
(Zhang and Zhong, 1980c). Sexual morphs and life cycle unknown.

Tinocallis (Pseudochromaphis) coreana Paik General colour not recorded

but the black head and prothorax with pale patches, the maculate forewings
and the pattern of small dark spots on the dorsal abdomen are probably very
distinctive in life; BL of alata 1.1-1.6 mm. On Hemiptelea davidii in China
(Zhang and Zhong, 1982c) and Korea (Paik, 1965). Sexual morphs and life
cycle unknown.

Tinocallis (Quednaucallis) distincta Ghosh, Ghosh and Raychaudhuri Alatae

pale (colour unrecorded), with dark antennae, tibiae, dorsal abdominal tuber-
cles, siphunculi and CAUDA; BL 1.8-2.2 mm. Described from an unidentified
leguminous plant in West Bengal (M.R. Ghosh et al., 1971); Chakrabarti
(1988) lists other records from West Bengal, mostly from undetermined plants,
but including one sample collected on Duabanga sonneratioides. Oviparae and
alate males in December-January, but alate viviparae are also present during
winter months so apparently in West Bengal there is only a partial holocycle.

Tinocallis (Sarucallis) himalayensis Ghosh, Ghosh and Raychaudhuri Alatae

greyish or greenish, somewhat waxy (J.H. Martin, pers. comm.); BL 1.2-
2.0mm. Forewings usually have dark spots at ends of veins and the branches
 Tinocallis 905

of the media are brown-bordered distally. Described fom an unidentified

leguminous plant (A.K. Ghosh et al., 1971e); specimens were subsequently
collected in India (West Bengal, Sikkim) on Peltophorum fleragineum
(Leguminosae) and Duabanga sonneratioides (Chakrabarti, 1988). Tinocallis
khonkaensis, described from undersides of leaves of Lagerstroemia macro-
carpa in Thailand (Danielsson and Robinson, 1978), seems to be a synonym,
as suggested by Ghosh and Quednau (1990); specimens from Lagerstroemia
sp(p). in East and Southeast Asia (Bangladesh, Thailand, Malaya, Singapore)
in the BMNH colln (leg. var.) confirm that there is no clear distinction
between these two species. Life cycle unstudied; alate viviparae and apter-
ous oviparae occur together in December-January in both northern India
and Thailand, indicating that there is only a partial holocycle. Males are
unrecorded.

Tinocallis insularis (Takahashi) All viviparae alate, greenish-yellow with

black-banded antennae; BL c. 1.6mm. On undersides of leaves of Sapindus
mukorossi in Taiwan (Takahashi, 1927a) and on Sapindus sp. in Japan
(Higuchi, 1972). Life cycle unknown. Also recorded from Pterocarya ste-
noptera (Tao, 1964), but these were probably vagrants. Tinocallis sophorae
Zhang, described from Sophora japonica in China (Zhang and Zhong, 1980c),
is similar and may be a synonym.

Tinocallis (Sarucallis) kahawaluokalani (Kirkaldy) Alatae broad-bodied,

pale yellow or yellow-green with dark brown markings (dark longitudinal
stripes on head and prothorax, dark brown pterothorax, transverse marks
on ABD TERG 1 and 2 incorporating the paired tubercles and distinctively
marked forewings); BL 1.2-1.8 mm. Immatures are greenish-yellow with pale
brown dorsal spots. On undersides of leaves of Lagerstroemia spp. and also
recorded from Lawsonia alba (Agarwala et al., 1989a). Widely distributed in
East and Southeast Asia; introduced to Italy, USA (where it is widely dis-
tributed, including Hawaii) and Puerto Rico. Oviparae and alate males in
September-October. Patti (1984) gave a concise general account with coloured
photographs. The ecology was studied in Italy by Patti et al. (1984), and in
North America by Alverson and Alien (1992). Mizeli and Knox (1993) studied
susceptibility of numerous Lagerstroemia cultivars in Florida. 2n = 6.

Tinocallis magnoliae Ghosh and Raychaudhuri Colour of alatae in life

unknown; BL c. 1.7mm. On Magnolia sp. in India (Meghalaya) (A.K.
Ghosh and Raychaudhuri, 1972b). Only collected once, possibly not on its
true host, but it does not fully agree with any known Tinocallis species on
Ulmaceae.

Tinocallis mushensis (Takahashi) Alatae pale yellow, with 3 longitudinal

brown stripes on head and pronotum; BL c. 2 mm. On young leaves of
Zelkova formosana in Taiwan (Takahashi, 1925). Life cycle unknown.
[Synonymized with zelkowae Takahashi, 1919b by Tao (1964), but on the basis
of the original description it seems to be a distinct species.]
906 Tinocallis

Tinocallis nevskyi Remaudière, Quednau and Heie Alatae pale yellow or

whitish, with very little pigmentation in spring and summer populations,
except for a black spot at the apex of the hind femur; BL 1.6-2.1mm.
On Ulmus spp. in Central and southwest Asia (Afghanistan, Iran, Turkey,
Georgia, Tadzhikistan, Pakistan, northern India); also on Zelkova crenata,
although Remaudière et al. (1988) thought that the occurrence of sexual
morphs on this plant in Iran was 'accidental'. Oviparae and alate males in
November. Remaudière et al. (1988) discussed differences from T. saltans.

Tinocallis (Quednaucallis) nigropunctata (Tao) Alatae yellowish-white with

banded antennae and a dark spot at base of pterostigma; BL 1.6-2.1 mm.
Described from an unidentified plant in China (Tao, 1964); subsequently
collected from Dalbergia hupeana (Zhang and Zhong, 1980c; as Tinocallis
dalbergiae). Some alatoid nymphs believed to be this species were collected
on Phyllanthus in Bhutan (A.K. Ghosh, 1976; as Sarucallis), but this is
unlikely to be a true host. Sexual morphs and life cycle unknown.

Tinocallis nikkoensis Higuchi Alatae pale yellow with pale or dusky anten-
nae and legs; BL c. 1.5mm. On Corylus spp. (heterophylla, sieboldiana) in
Japan (Higuchi, 1972). Biology and sexual morphs unknown. [Paik's (1965)
record of T. zelkowae on Corylus sieboldiana in Korea should perhaps be
referred to this species.]

Tinocallis platani (Kaltenbach), Plate l0b Alatae yellow to greenish-white

with extensive black-brown markings on head, thorax, abdomen and fore-
wings, and black siphunculi; BL 2.0-2.2 mm. On undersides of leaves of
Ulmus spp., especially U. laevis. Throughout Europe, across Asia to eastern
Siberia (Pashchenko, 1988b), and introduced into western North America,
where it occurs on U. americana. Oviparae and alate males in October.
Olkowski et al. (1982) reported biocontrol with an introduced parasitoid in
California.

Tinocallis saltans (Nevsky) Alatae yellow to orange-yellow, with brown

head and thorax even in early summer, with a more extensive distal black
patch on the hind femur and more forewing pigmentation than in nevskyi;
BL 1.4-2.2 mm (Nevsky, 1929b; Quednau, 1979). On Ulmus spp. in Spain
(Nunez Perez et al., 1991), Eastern Europe (Romania) and Central and East
Asia (eastern Iran, Tadzhikistan, Uzbekistan, Urals, Afghanistan, Siberia,
Korea and China (as yinchuanensis). Oviparae and alate males in October
(Nevsky, 1929b). Introduced into North America (Halbert and Pike, 1990).
[Accounts under this name from Western Europe (e.g. Heie, 1982) and
southwest Asia (Richards , 1967) should be referred to T. nevskyi (Remaudière
et al., 1988).] 2n = 16 (in China; Chen and Zhang, 1985b).

Tinocallis suzhouensis Zhang Alatae dusted all over with white powder; BL
c. 2.2cm. On Zelkova schneideriana in China (Zhang and Zhong, 1980c).
Sexual morphs and life cycle unknown.
 Tinocallis 907

Tinocallis takachihoensis Higuchi Alatae are pale yellow-green with shiny

black head and thorax, black distal section of hind femur and base of hind
tibia and black markings on the wings (Moritsu, 1983); BL 1.8-2.0 mm.
Recorded from Ulmus spp. in Japan (Higuchi, 1972), China and eastern
Siberia (Pashchenko, 1988b; as ussuriensis), and also from Hemiptelea davidii
in China (Zhang and Zhong, 1980c; as T. hemipteleae). Alatae have been
trapped and collected on Ulmus sp. in southern France (Quednau and
Shaposhnikov, 1988). Life cycle and sexual morphs unrecorded. 2n = 16*.
[Tinocallis sapporoensis Higuchi, described from three alatae on Ulmus sp.
in Japan (Higuchi, 1972), may be midsummer population of takachihoensis,
or possibly saltans; the absence of wing pigmentation in Higuchi's specimens
could be due to the method of slide preparation.]

Tinocallis (Sappocallis) ulmicola (Matsumura) Alatae greenish-white, with

black head and thorax, black apices of hind femora and bases of hind tibiae,
and 3 large black patches on each forewing; BL 1.4-1.6 mm. On undersides
of leaves of Ulmus spp. in Japan, China, Korea and eastern Siberia (Quednau
and Shaposhnikov, 1988). Life cycle and sexual morphs are apparently
unrecorded. 2n = 16.

Tinocallis ulmifolii (Monell) Alatae pale yellow to greenish, with white-

bordered dusky longitudinal streaks on head and pronotum; abdomen with
rows of white specks and dusky spots around hair-bases; BL 1.5-2.0 mm. On
undersides of leaves of Ulmus spp. (especially U. americana), widely distri-
buted in North America. Apterous oviparae and alate males in September-
October (Hottes and Frison, 1931; Palmer, 1952; as Myzocallis ulmifolii).

Tinocallis ulmiparvifoliae Matsumura Alatae pale bluish-green, developing

paired longitudinal white wax stripes on head and pronotum, a single, spinal
white stripe on pterothorax and white wax spots on dorsal abdomen (Moritsu,
1983). Tips of abdominal tubercles and distal ends of forewing veins are fre-
quently dark. BL 1.6-12.7 mm. On Ulmus parvifolia in Japan, Korea, China
and Taiwan. Introduced to Australia (Zeck, 1933, as Myzocallis viridis) and
England (on bonsai U. parvifolia; Stroyan, 1977). Oviparae recorded from
Korea (Paik, 1972). 2n = 16*. [Tinocallis viridis (Takahashi), on Zelkova in
Taiwan, is synonymized with ulmiparvifoliae by Tao (1958), but appears to
be a distinct species.]

Tinocallis viridis (Takahashi) Alatae green, with green eyes, banded anten-
nae, forewings with dark spots on pterostigma and at distal ends of veins,
dark abdominal tubercles and dusky brown siphunculi; BL 2.2-2.4 mm. On
Zelkova formosana in Taiwan. Life cycle and sexual morphs unknown. [This
species has been confused in the literature with ulmiparvifoliae.]

Tinocallis zelkovae Dzhibladze Alatae have dark brown head, thorax,

siphunculi and CAUDA, yellowish-green abdomen with extensive dark brown
paired patches or cross-bands and maculate forewings with extensive pigmen-
908 Tinocalloides

tation around veins; BL 1.4-1.8 mm. Described from Zelkova carpinifolia in

Georgia and also collected on Zelkova sp. in Iran (BMNH colln, leg. R. van
den Bosch). Monoecious holocyclic, with apterous oviparae and alate males
(Dzhibladze, 1957).

Tinocallis zelkowae (Takahashi) Alatae shining pale yellow or greenish-

yellow, with pale eyes, banded antennae, a fine brown mid-dorsal line on head
and pronotum, pale yellow legs and siphunculi and a pair of small brown
dorsal spots on each of ABD TERG 3-7; BL 1.4-1.8 mm. On undersides of
leaves of Zelkova serrata in Japan (Takahashi, 1919b), Korea and China, and
introduced to England on bonsai Zelkova (Stroyan, 1979; as T. nirecola).
Records from Ulmus may mostly be misidentifications of other species,
although this aphid does seem to be recorded frequently, as immature stages
as well as adults, on other plants; especially Betulaceae (Alnus, Carpinus) and
Leguminosae (Robinia, Glycine max and 'cultivated beans'). Monoecious
holocyclic in Japan, with alate males and apterous oviparae in October-
November; Inouye (1968) gives an account of biology and descriptions of all
morphs. 2n = 12*.

TINOCALLOIDES Basu Drepanosiphinae: Phyllaphidini

One species on Prunus in northern India and Nepal. A.K. Ghosh and Quednau
(1990) gave a full account.

Tinocalloides montanus Basu All viviparae alate, with brown head and
thorax, pale yellow abdomen with two widely-separated rows of black spino-
pleural patches; BL 2.1-2.6 mm. On undersides of leaves of Prunus spp.,
especially P. cerasus, in northern India (West Bengal, Himachal Pradesh) and
Nepal. Holocyclic, with apterous oviparae and alate males in December-
January. Sexuales occur on P. persica as well as on P. cerasus (Agarwala et al.,
1984). 2n = 18 (Kurl, 1981).

TOXOPTERA Koch Aphidinae: Aphidini

About five species resembling Aphis but with a stridulatory apparatus con-
sisting of ventrolateral ridges on the abdomen and peg-like hairs on the hind
tibia (Eastop, 1952). Of East Asian origin, with three species now widely
distributed on shrubs and trees. Martin (1991) provided a key.

Toxoptera aurantii (Boyer de Fonscolombe), Plate 13c, d Apterae oval,

shiny, reddish-brown or black, with banded antennae; BL 1.1-2.0 mm.
 Toxoptera 909

Immatures are brownish, alatae are brown-black with a black pterostigma

and, unusually for Aphidinae, a usually once-branched media. In dense,
ant-attended colonies on shoots and undersides of young leaves of host plants,
causing slight rolling, twisting or bending of mid-ribs. Large colonies produce
an audible scraping sound when disturbed. On many species of trees and
shrubs throughout warm temperate, subtropical and tropical regions (see also
B & E, 1984, p. 364). Starý et al. (1988) reported on biocontrol measures in
southern France, and Hussein and Kawar (1985) studied natural enemies
in Lebanon. 2n = 8. [Populations in which alatae have several secondary
rhinaria on ANT IV have been recorded on Celtis in Japan (as Aphis celtis
Shinji, 1922c) and on Ficus in China (as Toxoptera schlingeri). As alatae of
T. aurantii normally have only 0-1 rhinaria on ANT IV, these are possibly
a distinct species.]

Toxoptera citricidus (Kirkaldy) Apterae shining, very dark brown to black,

usually larger than aurantii, and with antennae less distinctly banded; BL
1.5-2.4 mm. Immatures are brown; alatae have a shiny black abdomen,
black ANT III, forewing with pale pterostigma and media 2-branched.
(Specimens in alcohol colour the fluid deep red.) In ant-attended colonies
on young growth of host plants, rolling leaves and stunting shoots. Mainly
on Rutaceae, especially Citrus, but occasionally large colonies develop on
young growth of other trees and shrubs (see also B & E, 1984, p. 365). In
southern Africa, Southeast Asia, Australia, New Zealand, Pacific Islands
and subtropical and warm temperate parts of South America. Apparently
anholocyclic everywhere except Japan, where a functional holocycle is now
reported (Komazaki, 1988). The large literature includes in recent years
laboratory studies of population parameters (Galatoire, 1983; Takanashi,
1989); field studies in Japan (Komazaki, 1988) and Venezuela (Rondón et al.,
1983 - including natural control by fungus and predators); and a review
of transmission of citrus tristeza virus (Roistacher and Bar-Joseph, 1987).
2n = 8.

Toxoptera odinae (van der Goot) Apterae grey-brown to reddish-brown; BL

1.3-2.4 mm. On undersides of leaves of host plants along main veins and
in dense colonies on young shoots, attended by ants. Rather polyphagous,
occurring on numerous shrubs and some trees in East and Southeast Asia and
recently becoming widespread in Africa south of the Sahara (Barbagallo and
Alcantara Santos, 1989; Martin, 1989). Apparently anholocyclic everywhere;
sexual morphs have not been recorded. [Martin (1991) described Toxoptera
victoriae from Zanthoxylum scandens in Hong Kong. This aphid may occur
on other Zanthoxylum that are of tree habit, and will key to odinae in the
polyphagous aphid key. Adults are, however, shiny black in life, only the
immatures being reddish-brown. For other distinguishing features see Martin
(1991).]

Toxoptera schlingeri Tao Apterae blackish-brown with banded antennae,

not or hardly distinguishable from T. aurantii; BL 1.6-1.8 mm. Described
from Ficus sp. in Hong Kong (Tao, 1961). There are alatae in the BMNH
910 Trichaitophorus

collection from Ficus microcarpa (leg. D.S. Hill) agreeing with Tao's descrip-
tion; also alatae with host unrecorded from China (Foochow; leg. M.S. Yang)
and one alata trapped in Nepal (leg. K.C. Sharma). Biology and life cycle
unknown.

TRICHAITOPHORUS Takahashi Chaitophorinae

Five East Asian species, at least four of them associated with Acer. Related
to Chaitophorus and Periphyllus, but with fused head and pronotum, and
with long thick, hollow hairs around margin of body, but not across dorsum.
Apterae have 5- or 6-segmented antennae. Chakrabarti and Mandal (1986)
revised the genus and keyed the species then known.

Trichaitophorus aceris Takahashi Apterae dark green with last two seg-
ments of antennae dusky, tarsi dark and siphunculi black (Takahashi, 1937a);
BL 1.4-1.8 mm. On young leaves of Acer spp.; originally described from
A. rubescens in Taiwan. Aphids identified as aceris have since been recorded
from various Acer spp. in northwest India and Nepal (Chakrabarti and
Mandal, 1986). However, there must be some doubt about the identity of the
Indian populations, as they are pale in life and have pale tarsi and siphunculi
(A.K. Ghosh, 1980). Alatae of Indian populations differ greatly from apterae,
having black-brown head and thorax, antennae and legs mainly dark, abdomen
with a large black patch on ABD TERG 4-6 and separate sclerites on
other tergites, and Chaitophorus-like siphunculi with polygonal reticula-
tion (Quednau and Chakrabarti, 1976; as Periphyllus pusillus). The popula-
tions identified as T. aceris in northwest India are monoecious holocyclic;
apterous males and oviparae were described by Chakrabati and Mandal
(1986), collected on Acer sp. in October-November.

Trichaitophorus aenigmatosus Pashchenko Apterae pale green to green; BL

c. 1.04mm. On Acer tegmentosum in eastern Siberia (Pashchenko, 1988b).
Other morphs and life cycle unknown.

Trichaitophorus japonicus Sorin Apterae green, with black-brown tarsi and

pale siphunculi; BL c. 1.6mm. On Acer tschonskii feeding on upper sides of
leaves along mid-rib and main veins (Sorin, 1979b). Other morphs and life
cycle unknown.

Trichaitophorus koyaensis Takahashi Apterae yellow with pale antennae,

legs and siphunculi; BL c. 1.2mm. On Acer rufinerve in Japan (Takahashi,
1961b). Other morphs and life cycle unknown.

Trichaitophorus recurvispinosus Hille Ris Lambers and Basu Apterae

flattened, pale glassy white with tips of antennae and legs dusky brown; BL
 Tuberaphis 911

2.4-2.5 mm. Stated to live on undersides of rolled leaves of a plant that was
unidentified at the time the description was published (Hille Ris Lambers and
Basu, 1966), but subsequently identified as Elaeocarpus sikkimensis (BMNH
colln, leg. DHRL). In West Bengal, India. Other morphs and life cycle
unknown. [The host plant and leaf-rolling habit is unlikely for a chaito-
phorine and needs additional confirmation. Probably the true hosts are Acer
spp., as listed by Chakrabarti and Mandal (1986) under T. aceris. However,
the size, colour in life, length of antennae and shape of marginal hairs make
it unlikely that the synonymy with T. aceris proposed by these authors is
correct.]

TSUGAPHIS Takahashi Hormaphidinae: ?Hormaphidini

One or two highly specialized conifer-feeding species of uncertain relationship

to other Hormaphidinae. Only apterous viviparae are known.

Tsugaphis sorini Takahashi Apterae pale greenish, including antennae and

legs; BL c. 0.9 mm. On undersides of leaves of Tsuga sieboldii in Japan
(Takahashi, 1957). Other morphs, biology and life cycle unknown.

Tsugaphis sorini piceicola Shaposhnikov and Gabrid Apterae pale green,

translucent, older specimens becoming dark brownish-green; BL c. 0.9 mm.
On Picea spp., especially pungens, schrenkiana, living in yellow depressions
on upper and undersides of needles, which eventually become twisted and
brittle, and drop prematurely. In Kirgizia. Anholocyclic, overwintering as
hibernating apterae (Shaposhnikov and Gabrid, 1987). Differences from sorini
s. str. are very small, and the two could be synonyms.

TUBERAPHIS Takahashi Hormaphidinae: Cerataphidini

About eight species are known in East and Southeast Asia, mostly described
from secondary hosts which are all Loranthaceae. Recently the link has been
made with primary host forms that produce elaborate branching galls on
Styrax, described in the genus Astegopteryx (Aoki and Kurosu, 1993).

Tuberaphis taiwana (Takahashi) Galls are large (about 14cm in diameter),

pale greenish-yellow and coral-like, with small circular distal openings, and
arise from the stem of Styrax formosanum in Taiwan (Fig. 124G, p. 577).
Alate emigrants (BL about 1.75mm) were collected in August (Takahashi,
1934b). The secondary host is unknown, but probably a member of the
912 Tuberculatus

Loranthaceae. Tao (1969; as Astegopteryx) synonymized this species with

A. vandermeermohri (Hille Ris Lambers), but this is unlikely because the gall
described for that species is quite different, and there are also differences in
the soldier morph (Aoki and Kurosu, 1993).

Tuberaphis takenouchii (Takahashi) The 'broccoli-head-like' gall of this

species on Styrax japonica (Fig. 1241) was redescribed by Aoki and Usuba,
(1989; as Aleurodaphis), who found that it contained sterile second instar
soldiers. S. Aoki (pers. comm.) has revised the generic position of this species
because of the type of symbionts it contains. Migration probably occurs to
a member of the Loranthaceae; Rappardiella plicator Noordam may be the
secondary host form of this aphid (S. Aoki, pers. comm.). For a description
of the alate emigrant see Takahashi (1934a; as Astegopteryx).

TUBERCULATUS Mordvilko Drepanosiphinae: Phyllaphidini

About 50 species of oak-feeding aphids with one or more tubercular abdom-

inal spinal processes. All viviparae and males are alate. The genus includes
some well-defined subgenera of limited distribution (Tuberculoides and
Camelaphis in western palaearctic; Orientuberculoides and Acanthocallis
in East Asia; Pacificallis in western nearctic; and Toltecallis in Mexico).
Richards (1968c) and Hille Ris Lambers (1974) reviewed the world fauna, and
accounts are available for Britain (Stroyan, 1977), northwest Europe (Heie,
1982), India (Chakrabarti, 1988; A.K. Ghosh and Quednau, 1990), China
(Zhang et al., 1990b), eastern Siberia (Pashchenko, 1988b) and Japan
(Higuchi, 1970). W. Zhang and G. Zhang (1991) did a cladistic analysis of
23 species in China. Quednau (1992) keyed the species of subgenus Pacificallis.

Tuberculatus (Tuberculoides) africanus Hille Ris Lambers Colour in life

unknown; BL 2.0-2.6 mm. Only known from Quercus mirbeckii (canariensis)
in Algeria (Hille Ris Lambers, 1974). Sexual morphs unknown. Closely related
to T. borealis.

Tuberculatus (Tuberculoides) albosiphonatus Hille Ris Lambers Alatae pale

green, with black-banded antennae, a sharply-bordered dark brown spot on
the pterostigma, and pale siphunculi; BL 1.1-1.7 mm (Hille Ris Lambers,
1974). On Quercus infectoria (incl. var. boissieri) in Iraq (Kurdistan) and
Israel (BMNH colln, leg. VFE). Sexual morphs unknown.

Tuberculatus (Tuberculoides) annulatus (Hartig) Alatae yellowish, greyish-

green or pink to purple in summer, with black-banded antennae, black tarsi,
and dark distal two thirds or more of siphunculi; BL 1.7-2.2 mm. On under-
sides of leaves of Quercus spp., especially Q. robur, less commonly Q. petraea.
 Tuberculatus 913

Throughout Europe, Siberia (Pashchenko, 1988b) and introduced on Euro-

pean oaks to Australia, New Zealand, North and South America. Alate males
and apterous oviparae occur in October (northern hemisphere). Rohitha and
Penman (1984) studied development under laboratory conditions, Kennedy
(1986b) studied the role of tarsi in host selection, Heimbach (1986) studied
population dynamics in relation to honeydew production, and Dahlsten et al.
(1990) reported specific parasitoids in Germany. 2n = 14.

Tuberculatus (Tuberculoides) borealis Krzywiec Alatae pale blue-green,

yellow mottled with green, or yellow, with blackish-banded antennae and
siphunculi only apically dark (rarely over more than distal half); BL 1.9-
2.3 mm. On Quercus robur, more rarely on Q. petraea or robur x petraea
hybrids, in Europe (UK, Belgium, Denmark, Norway, Sweden, Italy, Poland,
western Russia) and east to Iran (BMNH colln, leg. S.H. Hodjat). Sexual
morphs occur in October in northern Poland (Krzywiec, 1971).

Tuberculatus (Pacificallis) californicus (Baker) Alatae pale yellowish-green

(Baker, 1917a) with banded antennae and brown-black spots at bases of
all tibiae; BL 1.7-2.5 mm. On Quercus lobata in western USA (California,
Colorado, Oregon, Utah). Sexual morphs in November (BMNH colln, leg.
R. van den Bosch).

Tuberculatus (Orientuberculoides) capitatus (Essig and Kuwana) Alatae

pale green, greenish-yellow or whitish-yellow, antennae ringed with black, legs
pale; BL 2.4-2.6 mm. Under leaves of Quercus spp. (acutissima, aliena, fabri,
mongolica, serrata, variabilis) in East Asia (China, Taiwan, Korea, Japan).
Oviparae and males were illustrated by Moritsu (1983). 2n = 14. [A single
alata trapped in Korea was described as a subspecies, T. capitatus intermedius
(Hille Ris Lambers, 1974); Quednau (1979) redescribed this form from spe-
cimens collected in Korea on Castanea crenata.]

Tuberculatus (Pacificallis) chrysolepidis Quednau Colour of alatae in life

unknown; BL 1.9-2.9 mm. On Quercus chrysolepis in California, USA
(Quednau, 1992). Sexual morphs and life cycle unknown.

Tuberculatus (Pacificallis) columbiae Richards Colour in life not observed;

BL 2.0-2.5 mm. On Quercus garryana, recorded from Oregon (BMNH colln,
leg. DHRL) and British Columbia (Richards, 1965). Sexual morphs not
described.

Tuberculatus (Camelaphis) cornutus Richards Colour in life not recorded;

BL c. 1.9mm. On unidentified Quercus sp. in Turkey. Sexual morphs not
described, but oviparae were observed in October (Richards, 1969a).

Tuberculatus (Tuberculoides) eggleri Börner Alatae pale green or yellow-

green with orange thorax, banded antennae and siphunculi pale, or only dark
apically; BL 1.7-2.5 mm. On upper or undersides of leaves of Quercus
914 Tuberculatus

spp. (especially pubescens; also recorded from aegilops, cerris, petraea). In

southern, Central and Eastern Europe, eastward to Turkey and Moldavia.
Monoecious holocyclic on Q. pubescens, but it may overwinter as viviparae
in mild winters. Lampel (1974) discussed seasonal variation and differences
from T. borealis.

Tuberculatus (Tuberculoides) etruscus Barbagallo and Binazzi Alatae

yellowish-green, with head and thorax slightly brownish, and siphunculi darker
towards apices; BL 1.5-2.9 mm. On Quercus frainetto in Italy (Barbagallo
and Binazzi, 1991). Sexual morphs and life cycle unknown.

Tuberculatus (Orientuberculoides) fangi (Tseng and Tao) Alatae pale green

with banded antennae; BL 1.8-2.9 mm. On undersides of leaves of Quercus
acutissima in China, Korea and Japan (Hille Ris Lambers, 1974). Sexual
morphs not recorded.

Tuberculatus (Toltecallis) garciamartelli Remaudière and Quednau Alatae

whitish, with banded antennae and a pair of irregular pale green submarginal
stripes on the abdomen; BL 1.4-1.9 mm. On Quercus microphylla in Mexico
(Remaudière and Quednau, 1983). Sexual morphs unknown.

Tuberculatus (Orientuberculoides) grisipunctatus Zhang, Zhang and Zhong

Alatae green; BL 1.7-2.3 mm. On Quercus acutissima in Hebei Province,
China (Zhang et al., 1990b). T. pappus Zhang, Zhang and Zhong, described
from the same host and from Q. mongolica, is very similar if not a synonym.
Sexual morphs unknown.

Tuberculatus (Orientuberculoides) higuchii Hille Ris Lambers Alatae pale

yellow (Quednau and Shaposhnikov, 1988); BL 1.8-2.7 mm. On Quercus spp.
(dentata, mongolica, serrata) in Japan, Korea, China and eastern Siberia. In
Japan, populations with thicker and blunter abdominal spinal processes and
shorter PT were described by Hille Ris Lambers (1974) as a subspecies, T.
higuchii breviunguis, but might be seasonal variants. Sexual morphs unknown.

Tuberculatus (Acanthocallis) indicus L.K. Ghosh Alatae reddish to greenish-

yellow with pale eyes, banded antennae, mainly dark hind femora and bases
of hind tibiae and long black spinal processes on ABD TERG 2 and 3, con-
trasting with long pale processes on thorax and ABD TERG 1 (see Moritsu,
1983; as T. fulviabdominalis); BL 2.0-3.2 mm. On Quercus spp. (acutissima,
aliena, dentata, griffithii, mongolica, serrata) in India, Korea, China and
Japan. Life cycle unknown; alate viviparae were collected in December-
January in Manipur, India (Chakrabarti, 1988). This is the species referred
to in Japanese and Korean literature (e.g. Higuchi, 1972; Paik, 1972) as
T. fulviabdominalis (Shinji) - see Hille Ris Lambers (1974).

Tuberculatus (Tuberculoides) inferus Barbagallo Alatae yellowish-green,

lightly dusted with wax, with banded antennae and dark-tipped spinal pro-
 Tuberculatus 915

cesses on ABD TERG 3; BL 1.3-2.2 mm. Described from Quercus pubescens

(sensu lato) in Sicily, inhabiting only the undersides of the most basal leaves
on young plants (Barbagallo, 1990). Also known from Quercus sp. in Turkey
(BMNH colln, leg. DHRL). Life cycle and sexual morphs unknown.

Tuberculatus (Acanthocallis) japonicus Higuchi Alatae yellow to green or

brown, with pale eyes, mesothorax yellowish-green or brown dorsally and dark
brown ventrally, and dark green-black spinal processes on ADB TERG 2 and
3 (Quednau and Shaposhnikov, 1988); BL 2.5-2.9 mm. On Quercus spp. (den-
tata, mongolica), attended by ants, in Korea, Japan and eastern Siberia. Sexual
morphs were collected on Q. dentata (possibly the preferred host) in Korea in
November (BMNH colln, leg. W.H. Paik). Tuberculatus japonicus is one of
a complex of species including T. indicus; in China, a closely-related form with
characters somewhat intermediate between japonicus and indicus occurs on Q.
acutissima and Q. fabri, and was described as T. japonicus ssp. radisectuae by
Zhang et al. (1990b). 2n = 14 (for ssp. radisectuae; Chen and Zhang, 1985b).

Tuberculatus (Orientuberculoides) kashiwae (Matsumura) Alatae pale

yellow to pale greenish-yellow, sometimes yellowish-white or nearly white
(Quednau and Shaposhnikov, 1988); BL 1.4-1.9 mm. On undersides of leaves
of Quercus spp., especially Q. mongolica, in Japan, China, Korea and eastern
Siberia. Oviparae occur in Korea in early November (Paik, 1972). 2n = 14.
[Zhang et al. (1990b) described a species on Q. acutissima in China, T.
acuminatus, which will probably key to kashiwae, but the hairs on ANT III
and the front of the head are pointed, whereas those of kashiwae are blunt
or capitate.]

Tuberculatus (Pacificallis) kiowanicus Hottes Alatae are pale yellow,

yellowish-green or green, with head and thorax light brown, orange or pinkish,
often dusky on sides; BL 1.6-2.6 mm. On undersides of leaves of Quercus
gambelii and Q. gunnisonii in Colorado and Utah, USA. Sexual morphs were
collected on Q. gambelii in October (BMNH colln, leg. G.F. Knowlton) and
the male was described by Hottes (1949; as Myzocallis tonkawa). [Palmer's
(1952) account under the name M. maureri applies to this species.]

Tuberculatus (Orientuberculoides) konaracola (Shinji) Alatae pale green or

pale yellow-green to almost white, with banded antennae, brown marginal
stripes on pronotum, dark Cula and Cu1b in forewing, dark tibiae, brown-
black spinal processes on ABD TERG 3 and dark distal halves of siphunculi;
BL 1.7-2.8 mm. On Quercus serrata in Japan (Shinji, 1941); redescribed by
Hille Ris Lambers (1974) from alatae trapped in Korea. Holocyclic in Japan;
Shinji (1941) described the sexual morphs. Zhang et al. (1990b) described two
populations from Q. acutissima in China as subspecies of T. konaracola;
T. konaracola gansuensis and T. konaracola hangzhoensis.

Tuberculatus (Orientuberculoides) kunugi (Shinji) According to original

description, alatae are green with black eyes, banded antennae and with long
916 Tuberculatus

spinal processes on thorax and ABD TERG 1-3 that are all concolorous with
body (Shinji, 1924); BL not recorded. On Quercus acutissima and Q. serrata
in Japan (Higuchi and Miyazaki, 1969); no specimens are known, and it could
not be included in the key.

Tuberculatus (Camelaphis) maculipennis Hille Ris Lambers Alatae greyish

due to wax dusting, with pale antennae and legs, dark head and thorax,
blackish spinal and marginal processes, and maculate forewings; BL 1.0-
2.0mm. On Quercus spp. (infectoria, ?calliprinos, ?robur) in southwest
Asia (Turkey, Lebanon, Iraq) and also recorded from Q. pubescens in Italy
(BMNH colln, leg. S. Barbagallo). Sexual morphs in Turkey in October-
November (Hille Ris Lambers, 1974).

Tuberculatus (Pacificallis) maureri (Swain) Alatae light green to apple green

with red eyes, dusky-dark tibiae and forewing with a conspicuously dark-
bordered Cu1b vein; BL 1.4-2.2 mm. On Quercus agrifolia and Q. kellogii
in California, USA. Sexual morphs in early November (BMNH colln, leg.
DHRL).

Tuberculatus (Tuberculoides) maximus Hille Ris Lambers Colour in life

unknown, probably pale, with banded antennae, forewings with dark trian-
gular spots at distal ends of Cula and branches of media, and a rather dark
CAUDA; BL 2.0-3.1mm. On Quercus spp. (macranthera, persica) in
southwest Asia (Iran, Turkey, Armenia). Hille Ris Lambers (1974) discussed
seasonal and geographic variation; sexual morphs tentatively ascribed to this
species were collected on Q. macranthera in Iran in November.

Tuberculatus (Toltecallis) mexicanus Remaudière and Quednau Alatae very

pale green to whitish with a pair of dark lateral stripes on prothorax and a
pair of darker green lateral spots on each of ADB TERG 1, 3 and 6; BL
1.3-2.4 mm. On Quercus spp. (peduncularis, rugosa) in Mexico (Remaudière
and Quednau, 1983). Apparently anholocyclic.

Tuberculatus (Tuberculoides) moerickei Hille Ris Lambers, Plate 9b Alatae

pale yellow with banded antennae and distally dark siphunculi; BL 1.4-
2.6mm. On Quercus spp. (especially infectoria) in Mediterranean (Cyprus)
and southwest Asia (Iraq, Israel, Lebanon, Turkey). A sample from Q.
?pedunculate (= robur) in Turkey was described as a subspecies, T. moerickei
galatensis (Hille Ris Lambers, 1974). 2n = 14.

Tuberculatus naganoe (Shinji) According to original description, light green

with red eyes, banded antennae and black tibiae, and with pale finger-like
spinal processes on head, thorax and each abdominal tergite; BL c. 1.6mm.
On Quercus spp. in Japan (Shinji, 1941). No specimens are known and it could
not be included in the key on the basis of available information.
Tuberculatus (Tuberculoides) neglectus Krzywiec Alatae pale yellow, with
darker yellow mesothorax, eyes usually whitish-yellow, antennae banded,
 Tuberculatus 917

siphunculi dark except at base; BL 1.4-2.2 mm. On undersides of leaves

of Quercus petraea, more rarely on Q. robur or hybrids between the two.
In northern and northwest Europe (Britain, Ireland, Sweden, Denmark,
Germany, northern Poland). Sexual morphs in November (Krzywiec, 1965;
Heie, 1982).

Tuberculatus (Acanthocaltis) nervatus Chakrabarti and Raychaudhuri Col-

our of alata in life unrecorded; antennae banded, thorax probably dark with
postscutum black, forewing veins slightly bordered with fuscous, the long,
finger-like spinal processes on ABD TERG 1-3 are dusky to dark, and siphun-
culi are dark towards apices. BL c. 3.6mm. On unidentified Quercus sp(p).
in India (Meghalaya, Manipur, Nagaland: Chakrabarti and Raychaudhuri,
1976; Chakrabarti, 1988). Life cycle and sexual morphs unknown.

Tuberculatus (Orientuberculoides) paiki Hille Ris Lambers Alatae pale

greenish-yellow (Quednau and Shaposhnikov, 1988); BL 2.4-3.2mm. On
undersides of leaves of Quercus spp. (dentata, mongolica) in India (Megha-
laya), Korea, China and eastern Siberia. A.K. Ghosh and Quednau (1990)
provided a redescription. Sexual morphs and life cycle unknown. Regarded
as a subspecies of yokoyamai by Zhang et al. (1990b).

Tuberculatus (Camelaphis) pallescens Hille Ris Lambers Alatae bright

green, with banded antennae including short blackish-brown sections near
middle of ANT III; BL 2.5-2.5 mm (Hille Ris Lambers, 1974). On Quercus
spp. (calliprinos, infectoria, ?robur) in southwest Asia (Iraq, Lebanon,
Turkey), and also recorded from Q. pubescens in Sicily (Barbagallo and
Stroyan, 1982). Sexual morphs were collected on Q. calliprinos in Turkey in
November.

Tuberculatus (Pacificallls) pallidus (Davidson) Alatae pale green; BL 1.5-

2.2 mm. On leaves of a wide range of Quercus spp. (including non-native oaks)
in California. Partly anholocyclic; alatae were collected in December-January
in southern California. Separation from T. quercifolii, which is holocyclic on
native Californian oaks is difficult and needs further investigation.

Tuberculatus (Orientuberculoides) paranaracola Hille Ris Lambers Alatae

greenish-yellow or yellowish-green to pale green, with 'dark green reflecting
whitish' (Quednau and Shaposhnikov, 1988); BL 1.6-2.3 mm. On undersides
of leaves of Quercus mongolica (incl. var. grosseserrata) in Japan, China,
Korea and eastern Siberia. Hille Ris Lambers (1974) distinguished a sub-
species, T. paranaracola hemitrichus, from yellow traps in Korea, and this
subspecies is also recorded from China (Chen and Zhang, 1985b). Life cycle
and sexual morphs unknown. 2n = 14 (for ssp. hemitrichus).

Tuberculatus pasaniae (Davidson) Alatae narrow-bodied, pale green mottled

with darker green and with 'many delicate pruinose markings' (Davidson,
1915); eyes dark red, antennae ringed with black, spinal process whitish; BL
918 Tuberculatus

2.0-2.6 mm. On undersides of leaves of Lithocarpus densiflora in California,

USA. No sexual morphs known, and the occurrence of alate viviparae in
November to mid-February indicates at least partial anholocycly.

Tuberculatus (Pacificallis) passalus Quednau Colour of alatae in life

unknown; BL 1.3-2.2 mm. On Quercus dumosa in California USA. Oviparae
in late September-December, but apparently partially anholocyclic, as alate
viviparae occur in December and March (Quednau, 1992).

Tuberculatus (Acanthocallis) pilosus (Takahashi) Alatae yellowish-brown

with blackish eyes, and forewing veins with broad brown borders; BL
1.9-2.8 mm. Described from Quercus sp. in Taiwan (Takahashi, 1929) and
redescribed from Japan on Q. phillyraeoides (Higuchi, 1970). [There are some
discrepancies between the two descriptions which warrant further investiga-
tion.] Sexual morphs and life cycle unknown.

Tuberculatus querceus (Kaltenbach) Alatae dirty greenish to straw-coloured,

covered with fine powdery white wax, with a dark bifurcate spinal process,
a dark patch near end of hind femur and dark siphunculi; BL 1.4-2.4 mm.
On undersides of leaves of Quercus robur in Europe and east to Iran, Turkey
and the Caucasus (Heie, 1982). Sexuales in October.

Tuberculatus (Acanthocallis) quercicola (Matsumura) Alatae dark green or

mid-brown with blackish cross-bands on ABD TERG 1 and 2 bearing dark
(sometimes pale-tipped) processes and forewing veins dark-bordered; BL
1.8-2.9 mm. On upper and undersides of leaves of Quercus spp. (acutissima,
dentata, mongolica, variabilis), attended by ants. In Japan, Korea, China and
eastern Siberia. Quednau and Shaposhnikov (1988) noted some differences in
life between populations on Q. dentata and Q. mongolica. Sexual morphs and
life cycle not recorded. 2n = 16 in China and Japan (Blackman, 1986; Chen
and Zhang, 1985b).

Tuberculatus (Pacificallis) quercifolii (Davidson) Alatae pale green with

olive green head and mesothorax (Davidson, 1919); BL 1.2-2.4 mm. On leaves
of Quercus douglasii in California, USA. Apparently holocyclic; fundatrices
occur in April (BMNH colln, leg. DHRL), but sexual morphs have not been
recorded. Hille Ris Lambers (1974) probably had a mixture of T. quercifolii
and T. pallidus (see Quednau, 1992).

Tuberculatus (Orientuberculoides) querciformosanus (Takahashi) Alatae

yellow or yellowish-white with yellow eyes and antennae ringed with black; BL
1.5-2.5 mm. On Quercus spp. (dentata, mongolica, variabilis) in Japan, China,
Taiwan, Korea and eastern Siberia. Sexual morphs and life cycle unrecorded.

Tuberculatus (Tuberculoides) remaudierei Nieto Nafria Alatae pale green

with red eyes; BL 1.8-3.0 mm. On Quercus pyreniaca in Spain. Sexual morphs
in October (Nieto Nafria and Mier Durante, 1978).
 Tuberocephalus 919

Tuberculatus (Toltecallis) spiculatus Richards Alatae white to very pale

green; BL 1.9-2.4 mm. On Quercus rugosa in Mexico. A subspecies, T.
spiculatus rebecae, was described from the same host (Remaudi re and
Quednau, 1983). Sexual morphs unknown.

Tuberculatus (Acanthocallis) stigmatus (Matsumura) Alatae shining dark

brown to bluish-black with banded antennae, pale fore legs and black hind
legs (see Moritsu, 1983); BL 2.0-2.6 mm. Immatures pinkish--blue or pinkish-
brown with longitudinal rows of black spots. On Quercus spp. (acutissima,
aliena, dentata, mongolica, serrata, variabilis) in Japan, Korea, China and
eastern Siberia. Sexual morphs in October in Korea (Paik, 1972).

Tuberculatus (Neomyzocallis) tuberculatus (Richards) Alatae yellow with

variable dark pigmentation, wings clear or with diffuse fuscous pigmentation;
BL c. 1.5mm (Richards, 1965; as Myzocallis tuberculata). On Quercus spp.
(alba, macrocarpa, muehlenbergii, velutina) in northwestern USA and western
Canada. Sexual morphs and life cycle unrecorded. [N.B. Tuberculatus tuber-
culatus Richards, 1968c = Tuberculatus fangi.]

Tuberculatus yokoyamai Takahashi Alatae pale greenish-yellow or pale

yellow with whitish eyes and banded antennae (see Moritsu, 1983); BL 1.6-
2.4mm. On Quercus spp. (crispula, mongolica, serrata) in Japan, China,
Korea and eastern Siberia. Life cycle unknown. 2n = 14. [T. fuscotuber-
culatus, described from Q. acutissima in China (Zhang et al., 1990b) is similar
and may key to this species, but the spinal processes on ABD TERG 1 and
2 are dark brown (pale on 3) and a black spot on distal part of hind femur.]

TUBEROCEPHALUS Shinji Aphidinae: Macrosiphini

About ten species of Myzus-like aphids in East Asia, often with hair-bearing
siphunculi, galling the leaves of Prunus in spring. Where the life cycle is
known there is host alternation to Compositae (Artemisia, Gnaphalium). The
generations on secondary hosts look very different from the spring forms on
Prunus, and this has contributed to the taxonomie confusion in the group.
Accounts are available from Japan (Miyazaki, 1971), China (Chang and
Zhong, 1976) and eastern Russia (Pashchenko, 1988b). Moritsu and Tokumoto
(1972) gave an account of Japanese Tuberocephalus (under Myzus) on flower-
ing cherries in which they included several unnamed species. Sorin (1993) and
Remaudi re and Sorin (1993) described galls and life cycles of several species
that could not be included in our key to Prunus aphids.

Tuberocephalus higansakurae (Monzen) Spring colonies in galls caused by

upward rolling and thickening of edges of distal parts of leaves of cherry trees,
920 Tuberocephalus

the sausage-shaped leaf-roll becoming yellowish-green or red. Dark green

fundatrices give rise to reddish-yellow apterae (BL 1.5-1.9 mm) and alatae.
Alatae have a dusky yellow abdomen with a dark central dorsal patch and fly
to an unknown secondary host. Greenish-white apterae of a Tuberocephalus
that could be this species were collected on Gnaphalium (BMNH colln, leg.
RLB), but this needs experimental verification. Recorded from Prunus
cerasus, P. pseudocerasus, P. serrulata and P. subhirtella, and from Japan,
Korea and China; however, it seems that Chang and Zhong (1976) in China
may have had T. misakurae. Monzen (1929) recorded alatae (gynoparae)
returning to P. subhirtella in October-November, but did not describe the
sexual morphs.

Tuberocephalus jinxiensis Chang and Zhong Described from apterae only

(BL 1.4-1.8 mm), collected on Prunus humilis in China in June (Chang and
Zhong, 1976). Life cycle unknown.

Tuberocephalus liaongensis Chang and Zhong Apterae (BL 1.8-1.9 mm)

and alatae were collected from Prunus pseudocerasus in China in June (Chang
and Zhong, 1976). Life cycle unknown. 2n = 12 (Chen and Zhang, 1985a).

Tuberocephalus misakurae Moritsu and Hamasaki Spring colonies live in

sausage-shaped galls formed by rolling and thickening of edges of distal parts
of leaves of cherry trees, as in the closely-related T. higansakurae. Fundatrix
deep green or green; colour of apterous progeny not recorded, BL 1.5-2.0 mm.
Alatae have a yellowish-green abdomen with a dark central dorsal patch and
fly in May to an unknown secondary host. Recorded from P. pauciflora
and P. takenakae in Japan (Moritsu and Hamasaki, 1983) and from P.
pseudocerasus in China (BMNH colln, leg. VFE). 2n = 12*.

Tuberocephalus momonis (Matsumura) Spring colonies roll edges of leaves

of Prunus persica and turn them red. Apterae are yellowish-brown to dark
brown with a greenish tinge (Matsumura, 1917); BL 1.4-2.0 mm. Alatae have
a pale yellowish-brown abdomen without a distinct dorsal patch and are pro-
duced in June (BMNH colln, leg. VFE), but colonies persist into summer on
peach, and the life cycle is still unknown. In Japan, China, Korea and Taiwan.
This species has been confused in the literature with T. higansakurae, T.
misakurae and T. sakurae. The true momonis seems to be specific to P. persica
2n = 12 (Chen and Zhang, 1985a).

Tuberocephalus sakurae (Matsumura) Spring colonies cause terminal leaves

of Prunus (Cerasus) spp. to become extensively curled, twisted and yellowed
or reddened (see Moritsu, 1983). Apterae are deep green to black (fundatrices
paler), with black siphunculi; BL 1.5-1.8 mm. Immatures are bright greenish-
yellow. Recorded from P. cerasus, P. jamasakura, P. lannesiana, P. maximo-
wiczii, P. sachalinensis, P. triloba and P. yedoensis, and from Japan, China,
Korea and eastern Siberia (including Kuril islands; as Sorbaphis kurilensis
Ivanoskaya-Shubina, 1966). Alatae have dark dorsal abdominal cross-bands
 Tuberolachnus 921

only partially fused between segments. Life cycle is unknown. [N.B. Tao's
(1966) sakurae was probably Myzus yamatonis, and Miyazaki's (1971) descrip-
tion is partly of misakurae.]

Tuberocephalus sosakii (Matsumura) Conspicuous, brownish, elongate

pouch-like galls are produced on the upper sides of Prunus leaves between
two veins (see Moritsu, 1983). Recorded from P. jamasakura, P. lannesiana,
P. sachalinensis and P. serrulata, and from Japan, Korea and the Kuril islands
(Pashchenko, 1988b). Progeny of the fundatrix are all alate, with bright yellow
abdomen; BL 1.2-1.8 mm. They leave the gall in June through an opening on
the underside of the leaf and fly to found colonies on Artemisia princeps
(Monzen, 1929). Return migration occurs in October. Specimens collected on
Artemisia in western Malaysia (BMNH colln, leg. J.H. Martin) may also be
this species. [Tuberocephalus tianmushaensis Zhang, described from Prunus
sp. in China (Zhang and Zhong, 1980a), is apparently similar to T. sasakii
but has fundatrices with 5-segmented antennae and alatae with a black dorsal
abdominal patch.]

TUBEROLACHNUS Mordvilko Lachninae: Lachnini

Two species with a single large conical spinal process or tubercle on ABD
TERG 4, but otherwise not very closely related and placed in separate sub-
genera. A.K. Ghosh (1982b) provided an account including both species.

Tuberolachnus salignus (Gmelin), Plate 15c, d Apterae mid-brown to dark

brown with a large dark brown tubercle in the centre of the dorsum; BL
5.0-5.8 mm. Alatae have the forewing membrane unpigmented. On stems and
branches of numerous Salix spp. and also very occasionally recorded from
Populus. Large colonies build up in late summer and attract numerous
honeydew-feeding insects. Virtually cosmopolitan in distribution, wherever
willows grow or are planted, except Australasia. Apparently anholocyclic
everywhere; no sexual morphs are known. Mittler (1957, 1958a, b) used this
species in classic studies of aphid feeding and nutrition, and Hargreaves and
Llewellyn (1978) studied its ecological energetics. 2n = 20.

Tuberolachnus (Tuberolachniella) sclerata Hille Ris Lambers and Basu

Apterae blackish with a conspicuous reddish-brown to black broad-based
tubercles on the dorsum; BL 4.0-5.2 mm. In large colonies on undersides of
leaves, petioles and young shoots of Eriobotrya petiolata in West Bengal (at
2100 m); on leaves they form rows along the mid-rib and main veins (Hille Ris
Lambers and Basu, 1966). Specimens of this or a very similar species have also
been collected on E. japonica in Indonesia (BMNH colln, leg. D. Noordam).
Alatae and other morphs are undescribed, and life cycle is unknown.
922 Uichancoella

UICHANCOELLA Calilung Hormaphidinae: Nipponaphidini

One species in the Philippines and Indonesia on Lithocarpus, apparently

related to Schizoneuraphis. Only apterae are known.

Uichancoella gabrieli Calilung Apterae blackish-brown profusely covered

with white wax, leaving only centre of dorsum exposed; BL 1.2-1.6 mm. In
a large colony on twigs of Lithocarpus sp. at high altitude in Philippines
(Calilung, 1975) and Sulawesi (BMNH colln, leg. J.H. Martin). Other morphs
and life cycle unknown.

UROLEUCON Mordvilko (= Dactynotus Rafinesque)

Aphidinae: Macrosiphini

A large genus of medium-sized to rather large aphids (BL 2-4 mm) associated
almost entirely with Compositae and Campanulaceae. The very few records
from trees involve the two most polyphagous species, which occasionally occur
on plants in other families; U. ambrosiae (Thomas), a reddish-brown aphid
with black siphunculi and a pale CAUDA in North, South and Central
America, and U. compositae (Theobald), a shiny very dark red aphid with
siphunculi and CAUDA both black, distributed widely through the Old World
subtropical regions, and introduced to Brazil (see B & E, 1984, pp. 368-369).

UTAMPHOROPHORA Knowlton Aphidinae: Macrosiphini

About ten species of Myzus-like aphids with a wide range of host plant rela-
tionships (B & E, 1984, p. 321). The six North American species are mostly
associated with Rosaceae and/or Gramineae; one lives all year around on
Crataegus.

Utamphorophora crataegi (Monell) Apterae lemon to canary yellow, with 4

widely-spaced brown dorsal spots (see Johnson and Lyon, 1988); BL 1.7-
2.3 mm. Often forming large colonies on twigs, shoots and leaves of Crataegus
spp., causing curling when on leaves. Widely distributed in North America.
Monoecious holocyclic; oviparae and alate males in September-October
(Palmer, 1952).
 Wahlgreniella 923

VESICULAPHIS Del Guercio Aphidinae: Macrosiphini

About 12 species mostly associated with Ericaceae and/or Cyperaceae

(Miyazaki, 1980b). One species is described from Prunus, and is probably
misplaced in this genus; it may be an aberrant Myzus.

Vesiculaphis pruni Chakrabarti and Medda Appearance in life not recorded;

apterae broadly oval, sclerotized, BL 1.7-2.1 mm. Collected in May on Prunus
cornuta in Uttar Pradesh, India (Chakrabarti and Medda, 1989). Other
morphs and biology unknown.

WAHLGRENIELLA Hille Ris Lambers Aphidinae: Macrosiphini

About six species with elongate swollen siphunculi, associated mostly with
Rosa and/or Ericaceae. One species utilizes Arbutus as secondary hosts.

Wahlgreniella nervata (Gillette) Apterae are spindle-shaped, pale green with

dark tips to the long, slightly swollen siphunculi; BL 1.4-2.5 mm. Alatae have
a green abdomen with variably developed dark dorsal cross-bands, sometimes
coalesced into an irregular patch. Apparently heteroecious holocyclic in North
America between Rosa and Ericaceae, including Arbutus spp., although the
host alternation still awaits proper experimental verification. Introduced,
anholocyclic populations occur on both Rosa and Arbutus in Europe, but
are morphologically separable from each other and have therefore been
treated as separate subspecies; Arbutus-feeding populations in Europe are
usually referred to as W. nervata ssp. arbuti Davidson, following Hille Ris
Lambers (1949). However, as the situation has apparently arisen as a result
of separate, recent introductions to Europe from common North American
stock, a formal subspecific nomenclature seems inappropriate. {Wahlgreniella
nervata has also been recorded in North America from Heteromeles
(= Photinia) arbutifolia (Patch, 1938), but the host identification should
perhaps be treated with suspicion.]
924 Xenothoracaphis

XENOTHORACAPHIS Takahashi
Hormaphidinae: Nipponaphidini

One species on Quercus in Japan, apparently closely related to the type species"
of Thoracaphis (arboris), and perhaps not warranting a separate genus.

Xenothoracaphis kashifoliae (Uye) Apterae black, strongly sclerotized,

covered with powder; BL c. 1.5 mm. Alatae are black with blackish subcosta,
pterostigma and veins in forewing. On upper sides of leaves of Quercus glauca
and Q. acuta in Japan. Anholocyclic, with alatae appearing in November
(Takahashi, 1958b; as X. kashiwae).

YAMATOCALLIS Matsumura Drepanosiphinae: Drepanosiphini

Seven oriental species associated with Acer and related to North American
Drepanaphis, but with a subapical zone of reticulation on the siphunculi. All
viviparae are alate. Accounts are available from Japan (Higuchi, 1972, 1974)
and India (Chakrabarti, 1988; A.K. Ghosh and Quednau, 1990).

Yamatocallis acericola Higuchi Alatae green, with forewing membrane dif-

fusely pigmented towards wing-tip and siphunculi blackish-brown except for
pale bases; BL 4.1-4.3 mm. On upper surfaces of leaves of Acer carpinifolia
in Japan (Higuchi, 1974). Life cycle unknown.

Yamatocallis brevicauda Chakrabarti Colour of alatae in life not recorded,

wing membrane probably unpigmented and siphunculi black-tipped; BL
3.1-3.8 mm. On Acer villosum in Uttar Pradesh, India. Sexual morphs in
October (Chakrabarti, 1988).

Yamatocallis hirayamae Matsumura Alatae yellowish-brown, forewings

with a broad dark band of pigment traversing the anterior part of the
membrane from the base of Cula to the wing-tip; BL 2.6-3.5 mm. On Acer
spp. in Japan, China, Korea and eastern Siberia (Higuchi, 1972; Pashchenko,
1988b). Sexual morphs in early November (in Korea; Paik, 1972).

Yamatocallis obscura M.R. Ghosh, A.K. Ghosh and Raychaudhuri Alatae

green, forewings with membrane unpigmented, siphunculi dark brown distally;
BL 3.4-4.1 mm. On undersides of young leaves of unidentified plant (assumed
here to be an Acer sp.) in West Bengal. A.K. Ghosh and Quednau (1990)
provided a redescription. Life cycle unknown.
 Yezaphis 925

Yamatocallis sauteri (Takahashi) Alatae green with yellowish-brown head

and thorax, forewings with a broad band of pigment extending from ptero-
stigma and Rs along distal branches of media, and black-tipped siphunculi;
BL c. 4.5 mm. On an unidentified Acer sp. in Taiwan (Takahashi, 1927a). Life
cycle unknown.

Yamatocallis takagii (Takahashi) Alatae green, forewings with Rs and

distal branches of media thickly brown-bordered and blackish siphunculi;
BL 4.5-4.7 mm. On twigs of Acer mono (var. glabrum) in Japan (BMNH
colln, leg. RLB). Life cycle unknown. 2n = c. 48 (Blackman, 1986).

Yamatocallis tokyoensis (Takahashi) Alatae green, with most of forewing

membrane infuscated except for posterior margin, and siphunculi pale brown;
BL 2.0-2.9 mm. On leaves of Acer palmatum in Japan. Kan and Sasakawa
(1986) studied relations with a syrphid predator.

YAMATOCHAITOPHORUS Higuchi Chaitophorinae

One East Asian species on Acer, related to Trichaitophorus but with long,
thick spinal as well as marginal hairs, the latter being duplicated.

Yamatochaitophorus albus (Takahashi) Apterae elongate oval, whitish with

pale antennae and legs; BL 0.8-1.0 mm. Described from an unidentified Acer
sp. at high altitude in Japan (Takahashi, 1961b; as Trichaitophorus albus),
and also recorded from A. acuminatum ( = caudatum) in Uttar Pradesh, India
(BMNH colln, leg. S. Chakrabarti).

YEZAPHIS Matsumura Aphidinae: Macrosiphini (?)

One species described from Sasa in Japan and not subsequently identified. A
summary of the original description is provided here.

Yezaphis sasicola Matsumura Apterae broadly spindle-shaped, black; BL

c. 1.5 mm. Antennae 5-segmented, 1.2mm long, yellow except for ANT I and
II which are black, with ANT I gibbous on inner side. ANT PT/BASE almost
4.5. Pronotal tubercles small, yellowish, longer than broad. 'Legs black, tibiae
and tarsi yellow' (sic). Siphunculi cylindrical, short (0.1 mm) and very slender,
somewhat constricted at base, 'with some curved hairs'. 'CAUDA shorter than
siphunculi, but broader' (Matsumura, 1917).
 IV
TECHNIQUES

Here we give only a brief account of methods of collecting, preserving and

making slide preparations of aphids, with particular reference to the iden-
tification of species colonizing crop plants. For more detailed techniques of
aphid study consult van Emden (1972) or Minks and Harrewijn(1988).

COLLECTING METHODS
Aphids may be collected by sweeping or beating, or simply by careful search-
ing of appropriate parts of the plant. Beating onto a tray or card held under-
neath the branch may be useful for free-living aphids on leaves and shoots
of trees, especially Lachninae on conifers, but it is not very efficient for active,
winged aphids such as many Drepanosiphinae, adults of which will fly as soon
as disturbed leaving only the immatures and damaged adults to fall onto the
beating tray. Beating cannot be used to collect aphids that live in wax wool
such as Adelgidae, and is obviously useless for those making galls or feeding
on the trunk or large branches.
It is best, if at all possible, to examine the tree carefully and try to find
the aphids in situ, so that a representative sample of all available morphs and
developmental stages can be collected and information gained about the
feeding site and the size and appearance in life of the colony, if one is formed.
Aphids inhabiting the undersides of leaves can often be observed in silhouette
by looking up through a leaf against the sun. Ant-attended aphids, such as
Thelaxinae, Chaitophorinae and many Lachninae, can often be detected
by looking for ants moving up the trunk of the tree. Large numbers of
active flies and wasps may also indicate presence of honeydew-producing
Homoptera.
It is usually considered best to bring the aphids back to the laboratory
alive on a piece of the host plant, rather than to collect them directly into

926
 Techniques 927

preservative. Large polythene or glass tubes stoppered with cotton wool,

polystyrene sandwich boxes or polythene bags are suitable containers. A
piece of tissue paper should be included in each container to soak up excess
moisture, and plenty of air should be included in a polythene bag before it
is tied up. In warm weather it is preferable to place samples in an insu-
lated cool bag or box with ice blocks, or a large precooled vacuum flask,
especially if they are going to be left in a car for even a short time. The
reason for keeping specimens alive is that colonies often consist of mainly
immature individuals, and the proportion of adults in the sample can be
increased by keeping the aphids for a few days in a cool place before pre-
serving them. Adults, especially alatae, should be left for one to two days
after they have reached maturity in order to develop their full pigmenta-
tion. This also provides a method of rearing out any parasitoids. It is impor-
tant, of course, to make sure first that no predators are included in the
container!
Another method of collecting tree-living aphids is to look for their eggs
on the twigs, branches or trunk in late summer or early spring. Pick a sunny
day as many aphid eggs are shiny black and much more conspicuous in sun-
light. Any twigs found to have eggs on them can be cut off, brought back
to the laboratory and placed in water; the eggs are then likely to hatch quite
quickly, and the hatching fundatrices can often be reared to adult on the swell-
ing and breaking buds and young shoots, although to rear a second generation
fresh twigs will almost certainly be required.
It is preferable to note down the fullest possible collection data at the time
of collection, including host plant, locality and date, and it is also important
to note biological information such as the colour of the aphids in life (both
adults and immatures), feeding site, whether the aphids occur singly or form
a colony, and whether or not there is ant-attendance. It is probably best to
keep a field notebook for this purpose, and to give each sample a collection
number which is written on a slip of paper and inserted in the container with
the specimens. If the specific identity of the tree is unknown or uncertain then
leaves and, if possible, fruits should be collected for examination by a
botanist; notes should also be made on the form and texture of the bark, and
the growth habit and height of the tree.

PRESERVATION AND MOUNTING

Aphids for morphological examination should be preserved in tightly stop-
pered tubes filled with 80-90% ethanol. For prolonged storage one volume
of 75% w/w lactic acid may be added after a few days to every two volumes
of alcohol containing specimens, and the tubes plugged with cotton wool and
kept under alcohol or on a cushion of cotton wool soaked in alcohol, in an
air-tight glass jar.
Maceration to remove the soft tissues of the specimen prior to mounting
is best carried out with the specimen tubes in a water bath kept near boiling
point, or on a dry-block heater. The following stages are involved:
928 Techniques

1. Gently boil the specimens in 95% ethanol for 1-2 min.

2. Decant or pipette off ethanol, add about 1 cm depth of 10% potassium
hydroxide (KOH) solution, simmer for 3-5 min.
3. Decant or pipette off KOH solution and wash the specimens free of all
KOH using 5-6 changes of distilled or deionized water, leaving them to soak
for at least 5 min each time.
The water-based Berlese mountant has frequently been used in the past by
aphid workers, but balsam mounts are recommended because of their proven
permanence and resistance to a wide range of climatic conditions. The
macerated specimens need to be totally dehydrated and cleared before mount-
ing in Canada balsam. This can be done most simply using Martin's (1983)
method:
4. Remove distilled water, add 1 cm depth of glacial acetic acid and leave for
2-3 min. Pipette off and repeat with fresh glacial acetid acid. Pipette off.
5. Add clove oil as clearing agent (specimens will float). Leave for 10-20 min
until specimens are clear.
6. Transfer 1-2 aphids to a drop of fairly thin Canada balsam on a clean
microslide and quickly arrange them with body untwisted, dorsal side upper-
most and appendages spread out.
7. Dip a clean coverslip in xylene and immediately lower it carefully onto the
specimens displayed in the drop so as to spread the mountant evenly without
trapping air bubbles.
8. Dry the slide horizontally in an oven at 50°C for about 1 week.
Additional notes
• If the rostrum is short and the thorax is very dark, as is often the case in
alate specimens, try to displace the rostrum to one side so that the last
rostral segment (R IV+V) is not obscured by the thorax. On the other hand,
if the rostrum is long or the thorax pale then the rostrum is best placed
ventrally along the midline, so that its length and the shape of R IV+V can
be readily determined.
• Characters of the embryos contained within the maternal body, especially
the number, size and arrangement of dorsal hairs, are often used in species
identification, e.g. of alatae of Pemphiginae and Hormaphidinae emerging
from galls and of certain Drepanosiphinae. These characters can be observed
through the maternal cuticle in well-prepared specimens, but are better
displayed if a small incision is made in the abdomen and a few embryos
are carefully squeezed out into the mounting medium before putting on the
coverslip.

LABELLING AND STORAGE

Aphid workers have found that thick card labels, glued to the slide with
an impact adhesive, help to protect the coverslip and enable slides to be
stacked vertically while awaiting attention. It is advisable to standardize the
 Techniques 929

Fig. 135. Labelled slide preparation.

labelling; the system most frequently used is shown in Fig. 135. Slides may be
stored horizontally in trays, or vertically in slotted drawers or boxes. For
larger collections a compact and versatile system involves vertical storage of
slides in individual envelopes made of paper or cellulose acetate, with inter-
spersed tab cards providing the collection with an integral index (Eastop,
1985).
 v
BIBLIOGRAPHY

REGIONALLY CLASSIFIED WORKS ON TREE-DWELLING

APHIDS
In addition to the publications referred to in the text there are a number of works that describe the
forest aphid faunas of countries in Eastern Europe and southwest Asia: former Czechoslovakia -
Pasek, 1954; Baltic States - Rupais, 1969; Israel - Halperin et al., 1989; Kirgizia - Gabrid, 1989;
Latvian parks - Rupais, 1961; Turkey - Çanakçioglu, 1966; Ukraine - Mamontova, 1955; Russia -
Shaposhnikov, 1955 (includes Far Eastern species).

N.B. B & E (1984) = Blackman, R.L. and Eastop, V.F. (1984) Aphids on the World's Crops. Wiley,
Chichester.

LIST OF REFERENCES
Agarwala, B.K. (1982) (fundatrix and alata of Greenideoidea ceyloniae) Akitu 45, 1-2.
Agarwala, B.K. (1983) (notes on some aphids in Bhutan) Indian Agric. 27, 261-262.
Agarwala, B.K. (1988) (laboratory studies on development and fecundity of 3 aphid species) 4th Int.
Symp. Aphidology. Aphidological Society of India, pp. 19-20.
Agarwala, B.K. (1989) (biological notes on Cinara atrotibialis) Entomon (India) 14, 257-259.
Agarwala, B.K. and Dixon, A.F.G. (1986) (Cervaphis schouteniae) Indian Biologist 18, 37-39.
Agarwala, B.K. and Ghosh, A.K. (1984) (Indian check list) Rec. zool. Surv. India, Occ. Pap. No. 50,
71 pp.
Agarwala, B.K. and Mahapatra, S.K. (1990) (sexual morphs) Orient. Insects 24, 237-246.
Agarwala, B.K. and Raychaudhuri, D. (1982) (2 species of Cinara) Akitu No. 46, 4pp.
Agarwala, B.K. and Raychaudhuri, D.N. (1982) (male Periphyllus aesculi) Entomon (India) 7,
325-326.
Agarwala, B.K. and Roy, S. (1987) (Greenideoidea ceyloniae) Entomon (India) 12, 109-111.
Agarwala, B.K., Raychaudhuri, D. and Raychaudhuri, D.N. (1980) (parasites) Entomon (India) 5,
39-42.
Agarwala, B.K., Mondal, P.K. and Raychaudhuri, D.N. (1982) (new spp., Sikkim) Entomon (India)
7, 37-45.

930
 Bibliography 931

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Calilung, V.J. (1975) (Uichancoella gabrieli) Philipp. Ent. 2 (5), 322-325 (1973).
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David, S.K. (1959a) (South Indian aphids V) Indian J. Ent. 20, 124-129 (1958).
David, S.K. (1959b) (Astegopteryx formosana ssp.) Indian J. Ent. 20, 175-80 (1958).
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Halme, P.P. (1992) (Ephedrus koponeni) Ent. Fennica 3, 113-116.
Halperin, J. (1986) (Eulachnus rileyi in Israel) Phytoparasitica 14, 319.
Halperin, J., Binazzi, A. and Swirski, E. (1989) (aphids of forest and ornamental trees and shrubs
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Hampson, M.J. and Madge, D.S. (1986a, b) (Phorodon humuli morphometric variation) Agriculture,
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Hargreaves, C.E.M. and Llewellyn, M. (1978) (Tuberolachnus, ecology) J. anim. Ecol. 47, 605-613.
Harper, A.M. (1959a) (poplar gall aphids in Alberta I) Can. Ent. 91, 489-496.
Harper, A.M. (1959b) (poplar gall aphids in Alberta II) Can. Ent. 91, 680-685.
Harper, A.M. (1963) (Pemphigus betae, biology) Can. Ent. 95, 863-873.
Harper, A.M. (1966) (Pachypappa, Thecabius) Can. Ent. 98, 1212-1214.
Harper, A.M. and Macdonald, M.D. (1966) (chromosomes of Eriosomatinae) Can. J. Genet. Cytol.
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van Harten, A. (1979) (Paoliella browni capinganae) Agronomià lusit. 39, 85-96.
van Harten, A. and Coceano, P.G. (1981) (some aphids trapped in Udine Province, Italy) Boll. Lab.
Ent. agr. Filippo Silvestri 38, 29-51.
van Harten, A. and Ilharco, F.A. (1972) (Paoliella) Revta Cienc. biol. Lourenco Marques 5, 79-91.
Hashimoto, H. and Furuta, K. (1988) (P. californiensis) Jap. J. appl. Ent. Zool. 32, 169-175.
Hebert, P.D.N., Finston, T.L. and Foottit, R. (1991) (Melaphis rhois) Genome 34, 757-762.
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Heie, O.E. (1980) (The Aphidoidea of Fennoscandia and Denmark I) Fauna entomologica scand. 9,
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944 Bibliography

Heie, O.E. (1986) (The Aphidoidea of Fennoscandia and Denmark III. Pterocommatinae and
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Heie, O.E. (1992) (The Aphidoidea of Fennoscandia and Denmark IV. Aphidinae, Macrosiphini)
Fauna entomologica scand. 25, 188pp.
Heikinheimo, O. (1963) (new Finnish aphids II) Sworn, hyönt. Aikak. 29, 184-190.
Heikinheimo, O. (1966) (aphids caught in Norway) Norsk ent. Tidsskr. 13, 387-392.
Heikinheimo, O. (1984) (Cinara spp. in Finland) Notul. ent. 64, 33-49.
Heimbach, U. (1986) (E. tiliae, T. annulatus, honeydew) J. appl. Ent. 101, 396-413.
Heinze, K. (1960) (Central European Myzinae) Beitr. Ent. 10, 744-842.
Heinze, K. (1961a) (Central European Myzinae) Beitr. Ent. 11, 24-96.
Heinze, K. (1961b) (Central European Pterocommatinae) Z. angew. Zool. 1, 97-115.
Heinze, K. (1962) (European Lachnidae, Adelgidae, Phylloxeridae) Dt. ent. Z. 9, 143-227.
Heriot, D. (1938) (Eriosoma crataegi, E. lanigerum) Proc. ent. Soc. Br. Columb. 34, 22-32.
Hernandez, R. and Rodriguez, M. (1985) (pine aphid identification) Centra Agricola 12, 11-14.
Higuchi, H. (1968) (revision of Takecallis) Insecta matsum. 31, 25-33.
Higuchi, H. (1970) (revision of Japanese Tuberculatus) Insecta matsum. 32, 111-123.
Higuchi, H. (1972) (Japanese Callipterinae = Drepanosiphinae) Insecta matsum. 35, 19-126.
Higuchi, H. (1974) (Yamatocallis) Kontyû 42, 227-231.
Higuchi, H. and Miyazaki, M. (1969) (hosts of Japanese aphids) Insecta matsum. Suppl. 5, 1-66.
Hille Ris Lambers, D. (1931) (2 new Astegopteryx spp.) Miscnea zool, sumatr. 55, 1-5.
Hille Ris Lambers, D. (1933a) (Astegopteryx on Styrax) Miscnea zool. sumatr. 76, 1-4.
Hille Ris Lambers, D. (1933b) (Stomaphis mordvilkoi) Stylops 2, 197-201.
Hille Ris Lambers, D. (1945) (Sappaphis = Dysaphis devecta) Tijdschr. PlZiekt. 51, 57-72.
Hille Ris Lambers, D. (1947a) (Aphididae of Europe III) Temminckia 1, 179-319.
Hille Ris Lambers, D. (1947b) (some W European aphids) Zool. Meded. Leiden 28, 291-333.
Hille Ris Lambers, D. (1947c) (Periphyllus) Tijdschr. Ent. 88, 225-242 (1945).
Hille Ris Lambers, D. (1948) (on Palestine aphids) Trans R. ent. Soc. Lond. 99, 269-289.
Hille Ris Lambers, D. (1949) (Aphididae of Europe IV) Temminckia 8, 182-329.
Hille Ris Lambers, D. (1952) (aphid fauna of Greenland) Meddr Gr0nland 136, 1-33.
Hille Ris Lambers, D. (1953a) (Aphididae of Europe V) Temminckia 9, 1-176.
Hille Ris Lambers, D. (1953b) (aphids from Cocos nucifera, Astegopteryx host alternation) Agric.
J. Dep. Agric. Fiji 24, 211-215.
Hille Ris Lambers, D. (1954a) (some Eritrean aphids) Boll. Lab. Zool. gen. agr. Portici 33, 171-186
(publ. 1956).
Hille Ris Lambers, D. (1954b) (new Israel aphids) Bull. Res. Coun. Israel (B) 4, 276-283.
Hille Ris Lambers, D. (1956a) (revision of Cervaphis) Ent. Ber., Amst. 16, 130-136.
Hille Ris Lambers, D. (1956b) (Netherlands aphids) Tijdschr. Ent. 98, 229-249.
Hille Ris Lambers, D. (1956c) (lachnids from Elaeagnaceae) Z. angew. Ent. 39, 467-473.
Hille Ris Lambers, D. (1957) (Pistacia aphids) Bull. Res. Coun. Israel (B) 6, 170-175.
Hille Ris Lambers, D. (1960a) (N American Chaitophorus) Tijdschr. Ent. 103, 1-30.
Hille Ris Lambers, D. (1960b) (additional Greenland aphids) Meddr Grønland 159, 1-18.
Hille Ris Lambers, D. (1960c) (new genera and spp. from Canada) Can. Ent. 92, 251-265.
Hille Ris Lambers, D. (1962a) (new Toxopterella sp.) Fla Ent. 45, 143-148.
Hille Ris Lambers, D. (1962b) (4 new spp. from Canada) Can. Ent. 94, 1031-1039.
Hille Ris Lambers, D. (1965) (some Japanese aphids) Tijdschr. Ent. 108, 189-203.
Hille Ris Lambers, D. (1966a) (Italian aphids) Boll. Zool. agr. Bachic. 8, 1-32.
Hille Ris Lambers, D. (1966b) (some synonyms) Ent. Ber., Amst. 26, 124-126.
Hille Ris Lambers, D. (1966c) (aphids from Pakistan) Tidjschr. Ent. 109, 193-220.
Hille Ris Lambers, D. (1966d) (notes on California aphids) Hilgardia 37, 569-623.
Hille Ris Lambers, D. (1966e) (review of aphid polymorphism) A. Rev. Ent. 11, 47-78.
Hille Ris Lambers, D. (1967) (Neophyllaphis) Zool. Meded. Leiden 42, 55-66.
Hille Ris Lambers, D. (1968) (Neuquenaphis) Tijdschr. Ent. 111, 257-286.
Hille Ris Lambers, D. (1970a) (new Cavariella spp.) Mem. Soc. ent. ital. 48, 285-299 (1969).
Hille Ris Lambers, D. (1970b) (Tetraneura) Boll. Zool. agr. Bachic. 2, 21-101.
Hille Ris Lambers, D. (1970c) (Nearctaphis) Proc. K. ned. Akad. Wet. 73, 48-62, 63-74.
Hille Ris Lambers, D. (1971) (2 new Drepanosiphum spp.) Ent. Ber., Amst. 31, 72-79.
 Bibliography 945

Hille Ris Lambers, D. (1973a) (Masonaphis lambersi) Tijdschr. PlZiekt. 79, 159-161.
Hille Ris Lambers, D. (1973b) (notes on some oriental aphids) Orient. Insects 7, 239-258.
Hille Ris Lambers, D. (1973c) (Schizoneurata tissoti) Fla Ent. 56, 295-299.
Hille Ris Lambers, D. (1974) (Tuberculatus) Boll. Zool, agr. Bachic. 11, 19-82 (1972).
Hille Ris Lambers, D. and Basu, R.C. (1966) (some new and little-known Aphididae from India) Ent.
Ber., Amst. 26, 12-20, 27-36.
Hille Ris Lambers, D. and van den Bosch, R. (1966) (on the genus Hoplochaetaphis) Ann. Mag. nat.
Hist. 8, 561-564 (1965).
Hille Ris Lambers, D. and Hottes, F.C. (1962) (aphids from Alaska) Ent. Ber., Amst. 22, 112-120.
Hille Ris Lambers, D. and Rogerson, J.P. (1946) (M. padellus) Proc R.. ent. Soc. Lond. 15, 101-105.
Hille Ris Lambers, D. and Stroyan, H.L.G. (1959) (2 new spp., including Myzocallis schreiberi) Mem.
Soc. ent. ital. 38, 84-89.
Hille Ris Lambers, D. and Takahashi, R. (1959) (Thoracaphis, etc.) Tidjschr. Ent. 102, 1-16.
Hille Ris Lambers, D. and Wildbolz, T. (1958) (Allocotaphis quaestionis in Switzerland) Mitt.
Schweiz, ent. Ges. 31, 317-319.
Hintze-Podufal, C. and Thorns, H.J. (1979) (Chaitophorus leucomelas) Z. angew. Ent. 87, 388-392.
Hodjat, S.H. (1981) (Chaitophorus euphraticus) J. nat. Hist. 15, 365-374.
Hodjat, S.H. (1984) (aphids on Saliceae in Iran) Entomologie Phytopath. appl. 52, 5-6.
Hogmire, H.W., Brown, M.W., Schmitt, J.J. and Winfield, T.M. (1992) (population development,
insecticide susceptibility of A. pomi, A. spiraecola) J. ent. Sci. 27, 113-119.
Holman, J. (1974) Los áfidos de Cuba. Institute del Libro, La Habana, 304pp.
Holman, J. (1987) (Aphis of the Soviet Far East) Acta ent. bohemoslovaca 84, 353-387.
Holman, J. (1990) (new and little-known Aphis spp.) Acta ent. bohemoslovaca 87, 122-127.
Holman, J. and Szelegiewicz, H. (1972) (Mongolian aphids) Fragm. faun. 18, 1-22.
Holman, J. and Szelegiewicz, H. (1974a) (some Mongolian aphids) Annls zool. Warsz. 32, 1-17.
Holman, J. and Szelegiewicz, H. (1974b) (more Mongolian aphids) Acta ent. bohemoslovaca 68,
397-415.
Holman, J., Pelikan, J., Dixon, A.F.G. and Weismann, L. (eds) (1987) Population Structure,
Genetics and Taxonomy of Aphids and Thysanoptera. SPB Academic Publishing, The Hague,
542 pp.
Holopainen, J. and Heikinheimo, O. (1983) (Cinara maculata) Savon Luonto 7, 13-21.
Honda, H. (1921) (aphid spermatogenesis) Biol. Bull. mar. biol. Lab. Woods Hole 40, 349-369.
Hori, M. (1927) (new aphids from Hokkaido, Japan) Insecta matsum. 1, 188-201.
Hori, M. (1929) (new poplar aphids in Japan) Trans. Sapporo nat. Hist. Soc. 25, 109-115.
Hori, M. (1938) (Pemphigini in N Japan and Sakhalin) Insecta matsum. 12, 109-130.
Hottes, F.C. (1926) (2 new spp. from Minnesota) Proc. biol. Soc. Wash. 39, 111-114.
Hottes, F.C. (1933) (W Colorado aphids) Proc. biol. Soc. Wash. 46, 1-23.
Hottes, F.C. (1934) (aphid descriptions and notes) Proc. biol. Soc. Wash. 47, 1-8.
Hottes, F.C. (1949) (a new Myzocallis) Proc. biol. Soc. Wash. 62, 105-108.
Hottes, F.C. (1951) (Colorado juniper aphids) Proc. biol. Soc. Wash. 64, 145-146.
Hottes, F.C. (1952a) (Schizolachnus wahlea, Cinara spp.) Gt Basin Nat. 12, 57-61.
Hottes, F.C. (1952b) (Lachnus allegheniensis, etc.) J. Wash. Acad. Sci. 42, 127-129.
Hottes, F.C. (1952c) (Cinara wahhaka alata) Proc. biol. Soc. Wash. 65, 131-133.
Hottes, F.C. (1953a) (Cinara pinona, C. wahtolca) Proc. biol. Soc. Wash. 66, 153-158.
Hottes, F.C. (1953b) (Cinara wahhaka male) Proc. biol. Soc. Wash. 66, 199-202.
Hottes, F.C. (1954a) (alata of Lachnus alleghaniensis) Gt Basin Nat. 14, 21-22.
Hottes, F.C. (1954b) (sexual morphs of some Lachnini) Proc. biol. Soc. Wash. 67, 89-92.
Hottes, F.C. (1954c) (Cinara nimbata, C. puerca) Proc. biol. Soc. Wash. 67, 251-261.
Hottes, F.C. (1955a) (Cinara descriptions) Proc. biol. Soc. Wash. 68, 67-77.
Hottes, F.C. (1955b) (Cinara villosa, etc.) Proc. biol. Soc. Wash. 68, 101-104.
Hottes, F.C. (1955c) (3 new Cinara spp.) Proc. biol. Soc. Wash. 68, 197-203.
Hottes, F.C. (1956a) (Schizolachnus, key to spp. in USA) Proc. biol. Soc. Wash. 69, 59-62.
Hottes, F.C. (1956b) (Cinara acadiana) Proc. biol. Soc. Wash. 69, 63-64.
Hottes, F.C. (1956c) (Cinara soplada) Proc. biol. Soc. Wash. 69, 65-68.
Hottes, F.C. (1956d) (Cinara metalica, rustica) Proc. biol. Soc. Wash. 69, 83-88.
Hottes, F.C. (1956e) (Cinara sexual morphs, etc.) Proc. biol. Soc. Wash. 69, 89-92.
946 Bibliography

Hottes, F.C. (1956f) (Cinara grande, poketa, etc., and Schizolachnus flocculosa) Proc. biol. Soc.
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Hottes, F.C. (1956g) (Cinara bonica, C. bonita) Proc. biol. Soc. Wash. 69, 227-230.
Hottes, F.C. (1957a) (4 new Cinara spp.) Proc. biol. Soc. Wash. 70, 1-8.
Hottes, F.C. (1957b) (Cinara sexual morphs, etc.) Proc. biol. Soc. Wash. 70, 9-16.
Hottes, F.C. (1957c) (synopsis of Essigella) Proc. biol. Soc. Wash. 70, 69-109.
Hottes, F.C. (1958a) (new Cinara spp., etc.) Proc. biol. Soc. Wash. 71, 5-10.
Hottes, F.C. (1958b) (Cinara sitchensis) Proc. biol. Soc. Wash. 71, 61-62.
Hottes, F.C. (1958c) (Cinara nepticula) Proc. biol. Soc. Wash. 71, 63-64.
Hottes, F.C. (1958d) (Cinara contortae, C. parvicornis) Proc. biol. Soc. Wash. 71, 75-79.
Hottes, F.C. (1958e) (Cinara kuchea) Proc. biol. Soc. Wash. 71, 85-86.
Hottes, F.C. (1958f) (Essigella oregonensis) Proc. biol. Soc. Wash. 71, 155-156.
Hottes, F.C. (1958g) (Cinara jucunda) Proc. biol. Soc. Wash. 71, 171-172.
Hottes, F.C. (1958h) (Cinara harmonia) Proc. biol. Soc. Wash. 71, 191-195.
Hottes, F.C. (1959) (Schizolachnus lanosus) Proc. biol. Soc. Wash. 72, 13-14.
Hottes, F.C. (1960a) (Cinara wahsugae) Proc. biol. Soc. Wash. 73, 197-198.
Hottes, F.C. (1960b) (Cinara on Pinus edulis) Proc. biol. Soc. Wash. 73, 199-214.
Hottes, F.C. (1960c) (Cinara on Abies) Proc. biol. Soc. Wash. 73, 221-233.
Hottes, F.C. (1961a) (Cinara essigi) Proc. biol. Soc. Wash. 74, 1-2.
Hottes, F.C. (1961b) (Cinara on Picea) Gt Basin Nat. 21, 35-50.
Hottes, F.C. (1961c) (Cinara diabola, C. montanesa) Proc. biol. Soc. Wash. 74, 95-100.
Hottes, F.C. (1961d) (Cinara on Tsuga and Pseudotsuga) Proc. biol. Soc. Wash. 74, 111-118.
Hottes, F.C. (1964) (Cinara of Alaska) Ent. Ber., Amst. 24, 50-54.
Hottes, F.C. and Bradley, G.A. (1953) (Cinara canatra) Proc. biol. Soc. Wash. 66, 85-87.
Hottes, F.C. and Butler, G.D. (1955) (Cinara apacheca) Proc. biol. Soc. Wash. 68, 65-66.
Hottes, F.C. and Essig, E.O. (1953a) (2 new Cinara spp.) Proc. biol. Soc. Wash. 66, 159-172.
Hottes, F.C. and Essig, E.O. (1953b) (4 new Cinara spp.) Proc. biol. Soc. Wash. 66, 205-210.
Hottes, F.C. and Essig, E.O. (1954a) (Cinara hirsuta) Proc. biol. Soc. Wash. 67, 93-99.
Hottes, F.C. and Essig, E.O. (1954b) (Cinara azteca, dubia) Proc. biol. Soc. Wash. 67, 151-157.
Hottes, F.C. and Essig, E.O. (1955) (Cinara setulosa, etc.) Proc. biol. Soc. Wash. 68, 61-63.
Hottes, F.C. and Frison, T.H. (1931) (aphids of Illinois) Bull. Ill. nat. Hist. Surv. 19, 121-447.
Hottes, F.C. and Knowlton, G.F. (1954) (2 new Cinara spp.) Gt Basin Nat. 14, 11-13.
Hottes, F.C. and Wehrle, L.P. (1951) (Cinara tonaluca, etc.) Proc. biol. Soc. Wash. 64, 43-46.
Hottes, F.C., Essig, E.O. and Knowlton, G.F. (1954) (Schizolachnus) Proc. biol. Soc. Wash. 67,
273-274.
Howse, G.M. and Dimond, J.B. (1965) (Pineus pinifoliae on spruce) Can. Ent. 47, 952-961.
Hoy, M.A. and Cave, F.F. (1988) (resistant walnut aphid parasite) Calif. Agric. 42, 4-5.
Hrdy, L, Kremheller, H.T., Kuldova, J., Luders, W. and Sula, J. (1986) (insecticide resistance in
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Huang, X.S., Dai, J.X., Zhao, C.B. and Zhang, G.X. (1986) (Hyaloptems amygdali) Insect
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de Huiza, I.R. and Oritz, P.M.S. (1981) (Aphidiinae in Peru) Revta peru. Ent. 23, 129-132.
Hukusima, S. (1963) (Myzus malisuctus fecundity) Jap. J. appl. Ent. Zool. 7, 343-347.
Hull, L.A. and Starner, V.R. (1983) (D. plantaginea chemical control) J. econ. Ent. 76, 594-598.
Hussein, M.K. and Kawar, N.S. (1985) (T. aurantii natural control) Arab J. Pl. Protection 3, 11-17.
Iglisch, I. (1965) (Phylloxerina biology, morphology) Z. angew. Ent. 52, 325-474.
Ignoffo, C.M. and Granovsky, A.A. (1961a) (Mordvilkoja vagabunda) Ann. ent. Soc. Am. 54,
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Ignoffo, C.M. and Granovsky, A.A. (1961b) (M. vagabunda - gall) Ann. ent. Soc. Am. 54, 635-641.
Ilharco, F.A. (1966) (Thelaxes suberis) Agronomia lusit. 26, 83-89.
Ilharco, F.A. (1968) (Chaitophorus gomesi) Agronomia lusit. 29, 33-41.
Ilharco, F.A. (1971) (Paoliella hartem) Revta Cienc. Lourenco Marques 3(A), 35-43 (1970).
Ilharco, F.A. (1973) (Neophyllaphis viridis) Revt Cienc. Lourenco Marques 6(A), 29-35.
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Inouye, M. (1937) (Cinara spp. from Japan) Insecta matsum. 11, 100-105.
Inouye, M. (1938) (3 Lachninae spp. in N Japan and Sakhalin) Insecta matsum. 12, 74-80.
Inouye, M. (1939) (8 conifer aphids) Insecta matsum. 13, 132-142.
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Juchnevitch, L.A. and Kan, A.A. (1971) (Reclinasus buxtoni) Trudy Inst. Zool., AlmaAta 32, 12-17.
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 VI
INDEX TO SPECIES NAMES or APHIDS

abditus (-a, -um), Chaitophorus see aesculi, Pemphigus, Stomaphis yanonis

macrostychyae ssp.
abieticola, Cinara see confinis aethusae, Dysaphis crataegi ssp.
abieticolens, Pineus see pinifoliae affinis, Dysaphis, Pterocallis, Takecallis
abietihabitans, Cinara africanus, Sitobion, Tetraneura,
abietinus (-um), Elatobium, Mindarus, Tuberculatus
Pineus agilis, Eulachnus
abietis, Adelges, Cinara see agrifoliae, Atarsaphis, Neosymydobius •
matsumurana agrifolicola, Myzocallis
acadiana, Cinara ailanthi, Kaburagia
acanthopanaci, Aphis ajuscanus, Neosymydobius
acericola, Periphyllus, Yamatocallis akinire, Tetraneura see
acerifoliae, Drepanaphis nigriabdominalis
acerihabitans, Periphyllus alabastrum, Eriosoma see pyricola
acerinum, Drepanosiphum, Stomaphis alacra, Cinara
graffi ssp. alaskana, Cinara see hottesi
aceriphagus, Periphyllus alba, see albus
aceris, Drepanosiphoniella, albasiphus, Neosymydobius
Drepanosiphum, Neoprociphilus, albida, Pterocallis
Periphyllus, Stomaphis, albosiphonatus, Tuberculatus
Trichaitophorus albostriata, Pseudoregma see
actinodaphnis (-e), Aiceona bambusicola and dendrocalami
acuminatus, Tuberculatus see albus, Cinara see piceicola,
kashiwae Diphyllaphis, Gootiella,
acutihirsutus, Lachnus, Neobetulaphis, Trichaitophorus see
Mollitrichosiphum see nandii Yamatochaitophorus
acutirostris, Cinara alexanderi, Paraoregma see
aderuensis, Chaitoregma see tattakana Pseudoregma
adgnatum, Sitobion see phyllanthi allegheniensis, Lachnus
aegopodii, Cavariella allogenes, Periphyllus
aenigmatus, Adelges see laricis allozelkowae, Tinocallis
aequiunguis, Tetraneura almatinus, Brachycaudus

967
968 Index to Species Names of Aphids

alni, Boernerina, Calaphis, Illinoia, arctosetosus, Betulaphis see pelei

Mollitrichosiphum, Pterocallis, areolatus, Mexicallis
Stomaphis, Symydobius oblongus ssp. ariae, Dysaphis
alniarius, Symydobius aristolochiae, Paradoxaphis
alnicola, Eutrichosiphum, Hannabura, arizonica, Cinara
Mesocallis arma, Colophina
alnicolens, Betacallis armeniaca, Dysaphis
alnifoliae, Eutrichosiphum, armiger, Pineus see pinifoliae
Mollitrichosiphum see alni, artemisiae, Tüberocephalus
Pterocallis artocarpi, Greenidea
alnijaponicae, Pterocallis arunachali, Eutrichosiphum
alnosa, Calaphis arundicolens, Takecallis
alticola, Eulachnus see agilis arundinariae, Melanaphis,
alyeska, Essigella Neocranaphis, Rhopalosiphum,
amamiana, Nipponaphis see Takecallis
machilicola. . . .? asclepiadis, Aphis
ambrosiae, Uroleucon asiaticus, Myzus
amelanchericolens, Macrosiphum asiphon, Stomaphis
americanus, Eriosoma, Periphyllus, asiphum, Neopterocomma
Prociphilus, Symydobius aspidaphoides, Cavariella
amerinae, Plocamaphis assamense, Allotrichosiphum,
amygdali, Hyalopterus Eutrichosiphum, E. tattakanum ssp.,
amygdalinus, Brachycaudus, Myzus Metanipponaphis
analiliae, Mexicallis assetacea, Plocamaphis see flocculosa
andrhaka, Schoutedenia emblica ssp. asymmachia, Tetraneura
angelicae, Dysaphis aterrima, Paducia
anncharlotteae, Eriosoma atlantica, Cinara
annulatus, Tuberculatus atra, Cinara
anonae, Greenidea atrapinivora, Cinara see pinidensiflorae
antennata, Aleurodaphis, Monaphis, atripes, Cinara
Paducia atroalbipes, Cinara
anthrisci, Dysaphis atrotibialis, Cinara
antirrhinii, Myzus atrovirens, Taiwanaphis
anyangense, Pterocomma see yezoense atuberculata, Taiwanaphis
anzai, Cinara aucupariae, Dysaphis
apacheca, Cinara aurantii, Toxoptera
aphananthae, Stomaphis auratum, Eriosoma
apiifolia, Dysaphis aureus, Betulaphis see quadrituberculata
apini, Cinara aurlandicum, Acyrthosiphon
apulica, Cinara palaestinensis ssp. austriacus, Chaitophorus diversisetosus
aquatica, Aspidaphis see Cavariella ssp.
araliae, Cavariella autriquei, Sitobion
araucariae, Neophyllaphis autumna, Dinipponaphis
arboris, Thoracaphis see avellanae, Corylobium
Xenothoracaphis kashifoliae ayari, Paoliella
arbuti, Aphis, Wahlgreniella nervata ayyari, Greenidea see querciphaga
ssp. azaleae, Illinoia
arbutifoliae, Prociphilus caryae ssp. azteca, Cinara
archangelicae, Cavariella
archangelskii, Capitophorus baicalense, Pterocomma see
arctica, Calaphis groenlandica
 Index to Species Names of Aphids 969

baiccalensis, Paraprociphilus boerneri, Cinara (see cuneomaculata),

bailangense, Pterocomma Myzocallis, Pineus see pini
bakeri, Nearctaphis bogdanovi, Cinara see pruinosa
balachowskyi, Cinara bonica, Cinara
balsamiferae, Pemphigus see betae bonita, Cinara
balsamiferinus, Chaitophorus see borealis, Cavariella, Illinoia,
populifolii Pemphigus, Plocamaphis amerinae
bambucicola, see bambusicola ssp., Tuberculatus
bambusae, Astegopteryx, boycei, Pineus
Ceratogiyphina, Glyphinaphis, bozhkoae, Aphidura
Indoregma, Melanaphis brachychaeta, Pseudessigella
bambusicola, Chucallis, Myzocallis see brachycyclica, Dysaphis
Chucallis, Neocranaphis, brachysiphon, Plocamaphis flocculosa
Phyllaphoides, Pseudoregma, ssp.
Sitobion brachyunguis, Greenidea see kuwanai
bambusifoliae, Astegopteryx, Cerataphis bradleyi, Fullawaya
banksiana, Cinara braggii, Cinara, Drepanosiphum,
basalis, Astegopteryx, Calaphis see Essigella, Fullawaya
flava brancoi, Dysaphis
bathiaschvili, Roepkea marchali ssp. bratislavensis, Stomaphis
beccabungae, Aphis see frangulae brauni, Cinara
beibienkoi, Myzus braziliensis, Lizerius
bella, Myzocallis brevicauda, Yamatocallis
bengalensis, Ceratogiyphina, brevicornis, Pemphigus see populicaulis
Nippolachnus, Periphyllus brevipilosus, Betulaphis, Cinara,
berchemiae, Sitobion Eulachnus, Nippolachnus
berkemiae, Sitobion see berchemiae brevisetosus, Periphyllus, Subtakecallis
betae, Pemphigus, Smynthurodes brevispinosus, Cinara, Periphyllus
betulae, Euceraphis, Eutrichosiphum, breviunguis, Tuberculatus higuchii ssp.
Glyphina, Hormaphis, Stomaphis brideliae, Greenidea
betulaecolens, Calaphis brimblecombei, Neophyllaphis
betulaefoliae, Calaphis browni, Paoliella
betulaejaponicae, Euceraphis see bruneolineatus, Chaitophorus see
betulae ramicola
betulella, Calaphis buddleiae, Mollitrichosiphum see
betulicola, Calaphis nandii
betulinus, Hamamelistes bulbicauda, Neuquenaphis
beulahense, Pterocomma bulbosa, Fullawaya
beuthami, Chaitophorus see horii bulgaricus, Periphyllus
bhalukpongensis, Greenidea ceyloniae bumeliae, Prociphilus
ssp. bunii, Dysaphis see anthrisci
.bicolor, Pterocomma burrilli, Cinara
bipunctata, Phylloxera bursarius, Pemphigus
bisetosa, Monelliopsis burundiense, Sitobion
bisselli, Lachnochaitophorus see butzei, Neosymydobius
obscurus buxtoni, Rectinasus
biswasi, Cavariella
blumeae, Aleurodaphis see sinisalicis caerulescens, Euceraphis (see betulae),
bluncki, Eulachnus see rileyi Tetraneura
bodenheimeri, Myzocallis see caesium, Periphyllus
Hoplocallis pictus caliente, Cinara
970 Index to Species Names of Aphids

californica (-um, -us), Chromaphis, castaneae, Myzocallis, Phylloxera

Cinara, Essigella, Macrosiphum, castaneivora, Moritziella
Nearctaphis, Tuberculatus castaneoides, Myzocallis
californiensis, Periphyllus castanicola, Myzocallis
caliginosa, Aphis, Cinara see castanocallis, Castanocallis
coloradensis catalpae, Aphis
calligoni, Brachyunguis catalpinari, Shivaphis
calliptera, Callipterinella caucasica, Colopha, Drepanosiphoniella
calorai, Matsumuraja aceris ssp., Drepanosiphum, Dysaphis
calotropicus, Brachyunguis (see anthrisci), Stomaphis cupressi
calvus, Mexicallis, Neohormaphis ssp.
cambodiensis, Cervaphis see caudelli, Cinara
schoutedeniae cedri, Cinara
canadensis (-e), Adelges (see piceae), celastrii, Anuraphis see
Clydesmithia, Illinoia, Muscaphis, Ryoichitakahashia prunifoltae, Aphis
Neosymydobius (see albasiphus), celti, Shivaphis, Sumatraphis
Pseudopterocomma celticola, Shivaphis
canatra, Cinara celtis, Aphis see Toxoptera aurantii
canens, Chaitophorus see cembrae, Cinara, Eulachnus (see
macrostachyae pumilae), Pineus
cantonensis, Pseudoregma see cerasi, Myzus
bambusicola cerasicola, Brachycaudus see pilosus
capiangae, Paoliella browni ssp. cerasifoliae, Rhopalosiphum
capitatus, Tuberculatus cercidiphylli, Aulacorthum
capreae, Chaitophorus, Phylloxera see ceyloniae, Greenideoida
daphnoidis chaenomelis, Myzus see Ovatus
cardui, Brachycaudus malisuctus
Carolina, Cinara see atlantica chaerophylli, Dysaphis
carolinensis, Drepanaphis, Prociphilus chaerophyllina, Dysaphis
carpini, Greenidea, Myzocallis, chaetosiphon, Neobetulaphis,
Stomaphis Pterocomma, Sorbaphis
carpinicola, Greenidea, Kaltenbachiella, changaica, Tetraneura see yezoensis
Neochromaphis chengtuensis, Chuansicallis
carpinicolens, Macrosiphum cheni, Parathoracaphis, Prociphilus see
caryae, Longistigma, Monelliopsis, oriens
Prociphilus chetansapa, Aphis
caryaeavellana, Phylloxera chiangae, Paoliella
caryaecaulis, Phylloxera chibi, Cinara
caryaefallax, Phylloxera chilensis, Neuquenaphis
caryaefoliae, Melanocallis, Phylloxera chinensis, Astegopteryx,
caryaeglobuli, Phylloxera Schlechtendalia, Tetraneura
caryaegummosa, Phylloxera chinookiana, Cinara
caryaemagna, Phylloxera see choanotricha, Drepanaphis
caryaecaulis chomoensis, Pemphigus, see also
caryaeren, Phylloxera Epipemphigus imaicus
caryaescissa, Phylloxera chosoni, Lachnus see pallipes
caryaesemen, Phylloxera choui, Nurudea see meitanensis
caryaesepta, Phylloxera chrysolepidis, Tuberculatus
caryaevenae, Phylloxera chrysolepis, Neosymydobius
caryella, Monellia chuansicallis, Taoia
casimiri, Anomalaphis cicutae, Cavariella see salicicola
 Index to Species Names of Aphids 971

cimiciformis, Paracletus corylicola, Macrosiphum, Pterocallis

cinnamomiphila, Shivaphis see corylina, Illinoia
Machilaphis machili costata, Lachniella see Cinara
cinnamoniae, Euthoracaphis see covassii, Cinara
umbellulariae coweni, Adelges see cooleyi
circellatus, Pemphigus craccivora, Aphis
circumflexum, Aulacorthum crassicornis, Lachnus
cirsiiacanthoides, Aphis see fabae crataegarius, Ovatus
citricidus, Toxoptera crataegi, Dysaphis, Eriosoma,
citricola, Aphis (see spiraecola), Utamphorophora
Sinomegoura crataegicola, Prociphilus
cladogenous, Pineus crataegifoliae, Nearctaphis
clarus, Chaitophorus crataegistrobi, Prociphilus
clematis, Colophina crematogasteri, Dermaphis
clydesmithi, Nearctaphis cretacea, Eulachnus see agilis
cocciferina, Myzocallis crinitus, Chaitophorus
coccinea, Phylloxera cristafoliae, Hamamelistes
cognita, Cinara critchfieldi, Essigella
coloradensis, Calaphis, Cinara, Pineus cronartii, Cinara
columbiae, Tuberculatus crotonis, Eonaphis
comata, Cinara crucis, Chaitophorus
comes, Clethrobius cuneomaculata, Cinara
commatula, Cinara cupressi, Cinara, Siphonatrophia,
commensalis, Aphis Stomaphis
commiphorae, Paoliella curtihirsuta, Cinara
comperei, Anomalaphis see casimiri curtiwahtolca, Cinara wahtolca ssp.
compositae, Uroleucon curvipes, Cinara
compressa, Colopha curvispinosus, Schizolachnus
confinis, Cinara cuspidatae, Metanipponaphis
confusa, Phylloxera cylindricus, Pemphigus
congolense, Sitobion cynodonti, Asiphoniella
conica, Phylloxera cytisorum, Aphis
conifera, Phylloxera see depressa and
P. globosa form dactylidis, Forda see hirsuta
contortae, Cinara dactylonii, Asiphoniella
cooleyi, Adelges dahurica, Cinara
coracinus, Periphyllus dalbergiae, Tinocallis see
coreana, Plocamaphis, Thoracaphis see nigropunctata
Nipponaphis distyliicola, Tiliaphis, danesii, Foaiella
Tinocallis danubicus, Chaitophorus vitellinae ssp.
coreopsidis, Aphis daphnoidis, Phylloxerina
corni, Anoecia, Pseudasiphonaphis davidi, Eutrichosiphum
cornicola, Anoecia davidsoni, Phylloxera and also see
corniella, Aphis see salicariae Moritziella corticalis
cornifoliae, Aphis, Prociphilus, dearborni, Phloeomyzus see passerinii
Sitobion decaspermi, Greenidea, Taiwanaphis
cornutus, Myzus, Tuberculatus deducta, Euceraphis see lineata
corrugatans, Prociphilus delicatus, Chaitophorus see stevensis
cortecicolus, Pineus delottoi, Paoliella
corticalis, Moritziella dendrocalami, Pseudoregma
coryli, Myzocallis, Neochromaphis deplanata, Phylloxera
972 Index to Species Names of Aphids

depressa, Boernerina, Neothoracaphis eoessigi, Chaitophorus

(see Microunguis), Phylloxera eragrostidis, Geoica see utricularia
devastrix, Phylloxera eriobotryae, Hyalomyzus, Nippolachnus
devecta, Dysaphis see himalayensis
diabola, Cinara escherichi, Cinara see nuda
diacerivorus, Periphyllus see kuwanaii esfandiarii, Slavum
difficilis, Cinara see juniperivora essigi, Chaitophorus (see populifolii),
digitata, Cavariella Cinara, Essigella, Neuquenaphis,
dilaginosum, Eriosoma see patchiae Stegophylla
dipirivora, Sappaphis see piri etruscus, Tuberculatus
discolor, Myzocallis etsuhoe, Cinara
distincta, Tinocallis eugeniae, Aphis
distychii, Nipponaphis euonymi, Aphis
distylii, Reticulaphis euphorbiae, Macrosiphum
distyliicola, Nipponaphis euphraticus, Chaitophorus
divaricatae, Brachycaudus euscaphis, Sappaphis see Aphis horii
divaricatellus, Brachycaudus evodiae, Macrosiphum see
diversifolia, Chaitophorus see Acyrthosiphon
populialbae excelsae, Cinara see maritimae
diversis, Adelges see laricis exsiccator, Lachnus see pallipes
diversiseta, Cinara see hyperophila exultans, Myzocallis
diversisetosus, Chaitophorus ezoana, Cinara see pruinosa
dixoni, Drepanosiphum
donisthorpei, Paracletus fabae, Aphis
dorocolus, Chaitophorus, Pemphigus fagi, Phyllaphis, Platyaphis, Stomaphis
drepanosiphoides, Muscaphis fagicola, Mesocallis
dryobius, Clethrobius fagifoliae, Phyllaphis
dubium, Cinara, Eutrichosiphum, fangi, Tuberculatus
Neoacyrthosiphon, Pterocomma see farinosa, Aphis
populeum feminea, Aphis
durangaensis, Myzocallis fici, Greenidea, Reticulaphis distylii ssp.
ficicola, Greenidea, Nipponaphis
eastopi, Cinara, Essigella, Eumyzus filaginis, Pemphigus see populinigrae
echinata, Metanipponaphis, Paoliella fitchii, Prociphilus caryae ssp.,
edulis, Cinara Rhopalosiphum see insertum
edwardsi, Neuquenaphis flacourtiae, Greenidea
eggleri, Tuberculatus flava see flavum
ehretis, Rhopalosiphininus flavipes, Neuquenaphis see edwardsi
elaeagni, Capitophorus flavissima, Chaitophorus see capreae
elaeocarpi, Acyrthosiphon flavum, Astegopteryx (see basalis),
eligulatum, Eriosoma see auratum Calaphis, Dysaphis, Eriosoma,
elliotti, Myzocallis Eutrichosiphum, Oestlundiella
elongallis, Schlechtendalia flexilis, Cinara
elongata, Greenideoida, floccifera, Ceratovacuna
Mollitrichosiphum (see niitakaensis), flocculosus, Plocamaphis, Schizolachnus
Neothoracaphis, Parathoracaphis floccus, Pineus
emblica, Schoutedenia florentina, Phylloxera see querceus
engelmanni, Pineus see boycei floribundi, Myzocallis see
engelmanniensis, Cinara Globulicaudaphis pakistanicus
ensigallis, Macrorhinarium see floris, Carolinaia
Kaburagia rhusicola foaae, Phylloxera
 Index to Species Names of Aphids 973

foliodentata, Chaetogeoica ghanii, Pineus, Prociphilus

foliorum, Schizoneuraphis see gibberi, Hamamelistes
Distylaphis giganteus, Clethrobius (see comes),
follicularia, Forda see hirsuta Protopterocallis
formicaria, Forda gillettei, Euceraphis, Georgiaphis
formosanus, Aulacophoroides, gingerensis, Neophyllaphis
Cerataphis, Cinara, Cranaphis, ginnalae, Periphyllus see loricatus
Greenidea, Periphyllus, Prociphilus, glabra, Cinara, Kaltenbachiella,
Sinonipponaphis Phylloxera
fornacula, Cinara glacialis, Cinara
foveata, Phylloxera glandulae, Adelges
foveola, Phylloxera glandulosa, Astegopteryx, Myzocallis
foveolata, Reticulaphis distylii ssp. glaucae, Mollitrichosiphum (see
frangulae, Aphis nigrofasciatum), Neothoracaphis
fransseni, Cerataphis, Neophyllaphis glehna, Cinara
fraxini, Prociphilus globosa, Phylloxera
fraxinidipetalae, Prociphilus see globuli, Nipponaphis see Monzenia
americanus glyeines, Aphis
fraxinifolii, Prociphilus godavariense, Mollitrichosiphum
fresai, Cinara gombakana, Pseudoregma see
frisoni, Myzocallis pendleburyi
fritzmuelleri, Schizaphis see pyri gomboriense, Eriosoma see flavum
frosti, Grylloprociphilus see imbricator gomesi, Chaitophorus
fugans, Drepanosiphoniella aceris ssp. gooti, Parathoracaphis
fulviabdominalis, Anoecia, Tuberculatus gornitzi, Plocamaphis see flocculosa
see indicus ssp. brachysiphon
fulvicola, Indonipponaphis gossypii, Acyrthosiphon, Aphis
fumipenella, Protopterocallis gracilis, Cinara
funitecta, Adelges graffii, Stomaphis
furcatus, Chaitophorus, graminis, Colopha
Indiochaitophorus grande, Cinara
furcifera, Taiwanaphis grandifoliae, Phyllaphis
fusca, Essigella granovskyi, Calaphis (see flava),
fuscotuberculata, Tuberculatus Drepanaphis, Myzocallis
fusiformis, Tetraneura grassii, Crypturaphis
grisipunctatus, Tuberculatus
gabrieli, Uichancoella grobleri, Neophyllaphis
gairi, Pemphigus grodsinskyi, Hoplocallis see pictus
galatensis, Tuberculatus moerickei ssp. groenlandicus, Pemphigus (see
gallarum, Schizoneuraphis saliradicis), Pterocomma
gallica, Dysaphis see ariae grossa, Cinara see piceae
gallifoliae, Hormaphis see betulae grossae, Hamamelistes gibberi ssp.
gansuensis, Tuberculatus konaracola grossulariae, Eriosoma
ssp. guadarramae, Cinara
garciamartelli, Tuberculatus guangzhouensis, Greenidea see
garganicus, Cinara maghrebica ssp., formosana
Eulachnus see tuberculostemmata gyirongensis, Radisectaphis
garwhalensis, Eutrichosiphum,
Neothoracaphis, Periphyllus haldari, Greenidea
gentneri, Cinara, Fimbriaphis halli, Sitobion
georgiana, Phylloxera hamamelidis, Hormaphis
974 Index to Species Names of Aphids

hamiltoni, Macrosiphum hirticula, Cinara see moketa

hangnigri, Greenidea hispida, Monellia, Sinonipponaphis
hangzhouensis, Micromyzus, hissarica, Betulaphis
Neothoracaphis (see yanonis), hoerneri, Essigella
Shivaphis, Tuberculatus konaracolus hoffmanni, Ceratovacuna
ssp. hokkaidensis, Chaitophorus,
hannae, Greenideoida Periphyllus
harmalae, Brachyunguis horii, Aphis, Chaitophorus, Cinara
harmonia, Cinara hosoyai, Pineus
harpazi, Geoica hottesi, Cinara
harteni, Paoliella howardii, Glabromyzus see Carolinaia
harukawai, Pineus rhois
harunire, Eriosoma hughi, Pseudopterocomma
harvensis, Aspidophorodon humuli, Phorodon
hattorii, Cinara humulifoliae, Phorodon
havrylenkoi, Pineus see boerneri hyperophila, Cinara
heeri, Greenidea formosana ssp. hypogaeus, Chaitophorus
helianthi, Aphis hystrix, Paoliella
helichrysi, Brachycaudus
helvetica, Betulaphis see brevipilosa iaksuiense, Aphanostigma
hemipteleae, Tinocallis see ibofushi, Nurudea
takachihoensis ibotum, Aulacorthum
hemitrichus, Tuberculatus paranaracola idahoensis, Drepanaphis
ssp. ignotus, Pemphigus
henanse, Pterocomma see rufipes iliciphilus, Lachnus see pallipes
hendersoni, Cavariella ilicis, Aphis, Macchiatiella see
heraclei, Dysaphis crataegi ssp. Ryoichitakahashia prunifoltae,
heterophyllus, Pterocallis Phylloxera
heterotrichus, Euthoracaphis, imaicus, Pemphigus see Epipemphigus
Eutrichosiphum, Hoplochaitophorus imbricator, Grylloprociphilus
hidaense, Elatobium imbricatus, Pyrolachnus
higansakurae, Tuberocephalus immaculata, Astegopteryx,
higuchii, Tuberculatus Neobetulaphis
hillerislambersi, Cavariella (see immunis, Pemphigus
aquatica), Chaitophorus impatiens, Hydronaphis
himachali, Pseudothoracaphis incognita, Dysaphis
himalaica, Aiceona indicus, Avicennina, Cavariella,
himalayensis, Astegopteryx, Ceratovacuna, Chaitophorus, Cinara,
Capitophorus, Chaitophorus, Cranaphis (see formosana), Dysaphis
Chermes see Adelges knuchelii, pavlovskyana ssp., Kurisakia,
Nippolachnus see also Lachnus fici, Neotuberaphis, Parathoracaphisella,
Nipponaphis see Schizoneuraphis, Schizoneurella, Taoia, Tuberculatus
Periphyllus, Prociphilus, indigoferae, Anomalosiphon
Pseudastegopteryx see Astegopteryx inferus, Tuberculatus
minuta, Schizoneuraphis, Takecallis, inouyei, Chaitophorus
Tinocallis inscripta, Cinara
hippophaes, Capitophorus insertum, Rhopalosiphum
hirayamae, Yamatocallis insitus, Ovatus
hirsuta, Cinara, Forda insularia, Boernerina alni ssp.
hirsutustibus, Chromaphis insularis, Oregma see Astegopteryx,
hirticornis, Periphyllus Tinocallis
 Index to Species Names of Aphids 975

intermedia (-us), Cavariella, Cinara, karatanei, Nipponaphis

Eulachnus (see rileyi), Phylloxera, kashicola, Allotrichosiphum
Symydobius, Tuberculatus capitatus kashifoliae, Xenothoracaphis
ssp. kashiwae, Tuberculatus,
irae, Muscaphis drepanosiphoides ssp. Xenothoracaphis see kashifoliae
iranicum, Drepanosiphum kashmiricum, Eriosoma
isedakii, Adeiges see laricis kathleeni, Essigella
isensis, Metathoracaphis kaussarii, Forda
ishiharai, Adelges see pectinatae kayashimai, Parathoracaphis
iskanderkuli, Pemphigus keduensis, Ceratovacuna
israeliticus, Chaitophorus kenyensis, Paoliella see terminaliae
itadori, Macchiatiella kepongensis, Cerataphis fransseni ssp.
italica (-urn), Phylloxera, Pterocomma keshenae, Drepanaphis
iturbida, Myzocallis pepperi ssp. keteleeriae, Cinara
izas, Eutrichosiphum see pasaniae keteleerifoliae, Mindarus
khasyanum, Eutrichosphum
jacksoni, Pterocomma khonkaensis, Tinocallis see himalayensis
jamuritsu, Astegopteryx see basalis kiowanicus, Tuberculatus
japonensis, Dermaphis, Phorodon kirki, Essigella
japonicus, Adelges (see laricis), kitamiensis, Adelges see viridis
Aiceona, Betulaphis, Carolinaia, kiusa, Cinara
Cavariella, Ceratovacuna, Eriosoma, knowltoni, Chaitophorus (see
Kaltenbachiella, Lachnus (see macrostachyae), Drepanaphis,
tropicalis), Mindarus, Stomaphis, Essigella, Pemphigus
Trichaitophorus, Tuberculatus knucheli, Adelges
javanicus, Capitophorus hippophaes kochi, Cinara see kochiana
ssp., Mollitrichosiphum (see kochiana, Cinara
niitakaensis), Nipponaphis koelreuteriae, Periphyllus
jinxiensis, Tuberocephalus komareki, Myzocallis
joshii, Adelges konaracola, Tuberculatus
jucunda, Cinara konarae, Diphyllaphis
jugeshwari, Eutrichosiphum konoi, Cavariella, Cinara (see hattorii),
juglandicola, Chromaphis Prociphilus, Pterocomma see pilosa
juglandis, Callaphis see Panaphis konowashiyai, Pineus see pineoides
junctisensoriatus, Pemphigus koshuensis, Pseudoregma
juniperensis, Cinara koyaensis, Trichaitophorus
juniperi, Cinara, Sanbornia kozchukovae, Pterocomma
juniperivora, Cinara krahi, Sitobion
kuchea, Cinara
kabae, Symydobius kumaoni, Greenidea see nipponica
kachkouli, Aphis kunmingensis, Greenidea see okajimai
kagamii, Hamamelistes kunugi, Phylloxera, Tuberculatus
kahawaluokalani, Tinocallis kunzei, Dysaphis crataegi ssp.
kalipadi, Taiwanaphis kuricola, Myzocallis
kalopanacis, Aphis kurilensis, Sorbaphis see
kansugei, Colopha Tuberocephalus sakurae
kanzanensis, Greenidea see nigra kuwanai, Greenidea, Prociphilus
kanzensis, Drepanaphis kuwanaii, Periphyllus
kapuri, Chaitophorus
karafutonis, Adelges lachnirostris, Cinara
karamatsui, Adelges see laricis laciniatae, Eriosoma
976 Index to Species Names of Aphids

lactea, Pachypapella liukueinensis, Astegopteryx

laevis, Pineus see pini longicaudus, Aphis, Mexicallis spinifer
lambersi, Astegopteryx see Cerataphis ssp.
fransseni, Greenideoida, Illinoia, longicornus, Pemphigus
Pachypappa marsupialis ssp. longicornutum, Eriosoma
lanata, Neophyllaphis longifila, Ceratovacuna
lanigerum, Eriosoma longipennis, Cinara
lanosus, Schizolachnus see parvus . longipes, Chaitophorus
lantanae, Aphis longirostris, Cinara, Lachnus (see
lanuginosum, Eriosoma pallipes), Myzocallis, Paoliella,
laporteae, Hydronaphis Stomaphis
laportei, Cinara longirostrum, Lachnus see salicis
lapponicus, Adelges see laricis longiseta, Glyphina
lapponum, Chaitophorus longisetosus, Aiceona (see pseudosuga),
lariciatus, Adelges see laevis Chaitophorus, Greenidea (see
laricicola, Cinara see cuneomaculata querciphaga), Schizoneuraphis
laricifex, Cinara longiunguis, Chaitophorus, Myzocallis
laricifoliae, Cinara loricatus, Periphyllus
laricionis, Cinara lousianensis, Cinara
laricis, Adelges, Cinara, Elatobium, luchuanum, Mollitrichosiphum
Prociphilus lucifuga, Geoica
laserpitii, Dysaphis luminiferus, Betacallis
lataniae, Cerataphis lutea, Greenideoida, Schoutedenia see
lateralis, Brachycaudus see cardui ralumensis
lauberti, Dysaphis lutescens, Oregma see Astegopteryx
laurifoliae, Pemphigus bambusae
leeuweni, Astegopteryx see basalis luteus, Neosymydobius
lentiscoides, Slavum lyalli, Cinara
leonardi, Calaphis lycoposugus, Tiliphagus
leonidasi, Sitobion lyropictus, Periphyllus
leucomelas, Chaitophorus lythri, Myzus
lhasapopuleum, Pterocomma
liaongensis, Tuberocephalus machili, Machilaphis, Nipponaphis
lichtensteini, Pemphigus (see immunis), machilicola, Nipponaphis
Periphyllus see rhenanus machiliphaga, Nipponaphis (see also
ligustri, Myzus himalayaensis), Schizoneuraphis
ligustrifoliae, Prociphilus macrocanus, Pyrolachnus see pyri
lijiangensis, Cinara orientalis ssp., macrosiphum, Acyrthosiphon
Eutrichosiphum russellae ssp. macrostachyae, Chaitophorus
linderae, Aulacorthum maculata, Cinara see pilosa
lineata, Euceraphis maculatae, Aphis, Pterocallis
liquidambaris, Stomaphis maculipennis, Tuberculatus
liquidambarus, Longistigma maculipes, Cinara
liriodendri, Illinoia madagascariensis, Paulianaphis
lishanense, Eriosoma madronae, Aphis
lithocarpi, Astegopteryx (see maghrebica, Cinara
Lithoaphis), Mollitrichosiphum magnoliae, Aulacorthum, Neocalaphis,
lithocarpicola, Metanipponaphis Tinocallis
litseae, Aiceona (see titabarensis), magnolicolens, Neocalaphis
Eutrichosiphum, Nipponaphis maidis, Rhopalosiphum
litseicola, Schizoneuraphis majkopica, Dysaphis anthrisci ssp.
 Index to Species Names of Aphids 977

major, Anoecia melaina, Cinara

makabiae, Hamamelistes see betulinus melanocera, Myzocallis
makii, Eutrichosiphum melanosiphon, Chaitophorus
malaccensis, Oregma see Astegopteryx memecyloni, Taiwanaphis
basalis memorialis, Neosymydobius
malayana, Aiceona, Thoracaphis see mengluense, Eutrichosiphum see
Schizoneuraphis pasaniae
mali, Aphidounguis meridialis, Dysaphis see radicola
malicolens, Ovatus see crataegarius meridionalis, Myzocallis
malidauci, Dysaphis merkeri, Adelges
malisuctus, Myzus see Ovatus mensuae, Greenideoida see ceyloniae
malloti, Stomaphis metalica, Cinara
mammulata, Aphis mexicanus, Latgerina orizabaensis ssp.,
mamontovae, Periphyllus Muscaphis, Paraprociphilus,
manaliensis, Chaitophorus see indicus Tuberculatus
mandshuricus, Periphyllus michelbacheri, Neophyllaphis,
mangiferae, Greenidea Neuquenaphis
mangkamensis, Pemphigus micheliae, Formosaphis, Prociphilus
manipurense (-is), Eutrichosiphum, microgallis, Meitanaphis see
Hoplothoracaphis Schlechtendalia
manitobensis, Calaphis, Cinara, microsetosus, Hoplocallis, Monellia,
Sitobion see Macrosiphum Pemphigus
manoji, Eutrichosiphum (see microsiphon, Hoplocallis
quercifoliae), Nipponaphis microtrema, Diphyllaphis
maoi, Sinochaitophorus mimeuri, Geoica
marchali, Roepkea mimicus, Eriosoma, Myzocallis,
marginalis, Epipemphigus Neosymydobius
marginata, Forda, Pemphigella see mimosae, Sitobion see takahashii
Forda sichangensis minima, Aphis, Phylloxera, Prociphilus
mariana, Cinara see glehna see piceaerubensis
maritimae, Cinara minoripinihabitans, Cinara
marsupialis, Pachypappa minutissima, Callipterinella
martini, Plocamaphis minutus, Astegopteryx (see basalis),
matileae, Pterasthenia Chaitophorus, Cinara, Periphyllus,
matsumurai, Chaitophorus, Pemphigus, Shenawheum, Symydobius
Pineus mirabilis, Sinocallis (see Dasyaphis
matsumurana, Cinara rhusae), Reticulaphis
maui, Cinara see fresai misakurae, Tuberocephalus
maureri, Tuberculatus miscanthi, Sitobion see bambusicola
maximus, Tuberculatus miyabei, Hamamelistes
maxsoni, Georgiaphis see Eriosoma miyazakii, Chaitophorus
mimicum moerickei, Tuberculatus
medina, Monellia moketa, Cinara
mediocornutum, Eriosoma see ulmi momii, Elatobium
medispinosa, Cinara momonis, Tuberocephalus
mediterraneus, Cinara (see tujafilina), monelli, Chaitophorus, Drepanaphis
Eulachnus see tuberculostemmata, mongolicus, Chaitophorus nigricantis
Myzocallis ssp., Cinara, Pemphigus
meghalayensis, Capitophorus, monophagus, Pemphigus
Melanaphis monotuberculata, Paoliella see browni
meitanensis, Nurudea montana see montanus
978 Index to Species Names of Aphids

montanensis, Cinara neimongolense, Pterocomma see

montanesa, Cinara yezoense
montanicola, Cinara see pini, nekoashi, Ceratovacuna
Taiwanaphis neoalnicola, Eutrichosiphum see
montanus, Lachnus (see alleghenensis), tattakanum
Pemphigus (see matsumurai), neobetulella, Calaphis
Periphyllus, Pineus (see pinifoliae), neoficicola, Greenidea
Pseudoregma, Pterocallis, neogillettei, Aphis
Tinocalloides nepalensis, Callaphis see Panaphis,
monzeni, Nipponaphis Sinomegoura
mordvilkoi, Chaitophorus, Cinara, nepticula, Cinara
Diphyllaphis, Dysaphis, Forda (see nerii, Aphis
hirsuta), Pemphigus, nervatus, Tuberculatus, Wahlgreniella
Sinochaitophorus, Slavum neubergi, Cinara
moriokaensis, Colopha see Paracolopha nevskyi, Periphyllus, Tinocallis
morrisoni newelli, Cinara
moriokense, Eriosoma nicotianae, Myzus
morrisoni, Illinoia, Paracolopha niger (nigra, -um), Aphis, Carolinaia,
mucida, Euceraphis Cavariella, Chaitophorus, Cinara,
mugnosae, Stegophylla Greenidea, Juncomyzus see
mukerjee, Eutrichosiphum see Carolinaia, Phyllaphis,
nungsireiae Rhopalosiphum
multisetosa, Dysaphis (see pyri), nigeriense, Sitobion
Myzocallis, Nipponaphis nigra see niger
mumecola, Myzus nigrae, Chaitophorus
muradachi, Aulacorthum nigratibialis, Aphis
murphyi, Anomalosiphum nigrescens, Nearctaphis californica ssp.
murrayanae, Cinara nigriabdominalis, Mollitrichosiphum,
musae, Rhopalosiphum Tetraneura
mushaensis, Myzus nigricans, Drepanaphis
mushana, Greenidea nigricantis, Chaitophorus
mushensis, Tinocallis nigricentrus, Chaitophorus
muticae, Geoica see utricularia nigricola, Eulachnus
myersi, Aloephagus nigripes, Cinara
myricae, Calaphis, Greenidea, nigritus, Chaitophorus, Cinara
Myzocallis nigrofasciatum, Mollitrichosiphum
nigropunctata, Monelliopsis, Tinocallis
nachensis, Paoliella nigrostriata, Pterocallis
naganoe, Tuberculatus nigrum see niger
nainitalensis, Pemphigus niisimae, Epipemphigus
namaachae, Paoliella niitakaensis, Mollitrichosiphum
nanae, Myzocallis nikkoensis, Tinocallis
nandii, Mollitrichosiphum nimbata, Cinara
napaeus, Pemphigus nipponica, Cavariella, Greenidea
narafoliae, Eutrichosiphum see sinensis nirecola, Chromocallis, Kaltenbachiella,
nassonowi, Chaitophorus Tinocallis see zelkowae
nasturtii, Aphis nirmalae, Paoliella
nebrodensis, Adelges see piceae nishiyae, Watabura see Aphidounguis
neelagiriensis, Astegopteryx see minuta mali
neglectus, Chaitophorus, Tuberculatus nishyae, Dryopeia see Paracolopha
negundinis, Periphyllus morrisoni
 Index to Species Names of Aphids 979

nitidula, Cinara osmanthae, Prociphilus

nittakayamensis, Sinolachnus ostryae, Pterocallis
niuginii, Taiwanaphis osugiensis, Periphyllus viridis ssp.
nobukii, Rhopalosiphoninus osugii, Aiceona see malayana
nodulosus, Chaitophorus ovatirhusicola, Kaburagia
nordmannianae, Adelges ovogallis, Macrorhinarium see
nortonii, Pemphigus Kaburagia rhusicola
notabilis, Phylloxera
nothofagi, Taiwanaphis pacifica, Cinara
nudus, Chaitophorus, Cinara padellus, Myzus
nungsireiae, Eutrichosiphum padi, Rhopalosiphum
nusslini, Adelges see nordmannianae pahanensis, Melanaphis
nymphaeae, Rhopalosiphum paiki, Tetraneura, Tuberculatus
nyssae, Phylloxerina pakansus, Capitophorus
pakistanicus, Chaitophorus,
obesinymphae, Pemphigus see Globulicaudaphis
populitransversus palaestinensis, Cinara
obliquus, Mindarus pallescens, Tuberculatus
oblongus, Cinara (see pini), palliceps, Neuquenaphis
Symydobius pallidipes, Chaitophorus, Cinara see
obscurus, Cinara, Lachnochaitophorus, obscura
Megophyllaphis, Periphyllus, pallidus, Drepanaphis (see simpsoni),
Schizolachnus (see pineti), Dysaphis crataegi ssp.,
Yamatocallis Kaltenbachiella, Oregma see
obtusirostris, Mesocallis Astegopteryx basalis, Periphyllus,
occidentalis, Adelges (see piceae), Tuberculatus
Boernerina, Cinara, Myzocallis, pallipes, Lachnus
Nearctaphis crataegifoliae ssp. palmae, Cerataphis see fransseni
occulta, Myzocallis palmerae, Cinara see pruinosa
ochnearum, Sitobion pandani, Astegopteryx
ocoteae, Lizerius papillata, Neuquenaphis (see edwardsi),
odaiensis, Betacallis Paoliella, Sitobion
odinae, Toxoptera pappus, Tuberculatus see grisipunctatus
oenotherae, Anoecia paranaracolus, Tuberculatus
okajimai, Greenidea parasiticum, Eriosoma yangi ssp.
oleae, Prociphilus parasorbi, Dysaphis
onigurumi, Dasyaphis (see rhusae), parvicornis, Aiceona, Cinara
Kurisakia parvula, Hoplochaetaphis see
ontakensis, Euceraphis zachvatkini
ontarioensis, Cinara, Fullawaya parvus, Chaitophorus, Drepanaphis,
oregonensis, Adelges (see laricis), Neosensoriatus, Schizolachnus
Cinara, Drepanosiphum pasaniae, Eutrichosiphum, Tuberculatus
oregoni, Cinara pasheki, Cinara see maghrebica
oriens, Prociphilus passalus, Tuberculatus
orientalis, Cinara, Forda, Patchiella passeki, Pemphigus
reaumuri ssp., Pineus, Prociphilus passerinii, Phloeomyzus
(see kuwanai), Prociphilus xylostei pastinacae, Cavariella
ssp. (see ussuricus), Schizolachnus patchae, Chaitophorus populicola ssp.,
orizabaensis, Latgerina Pineus see similis
ornatus, Myzus patchiae, Eriosoma
osborni, Cinara patriciae, Illinoia
980 Index to Species Names of Aphids

pavlovskyana, Dysaphis pini, Cinara, Essigella, Eulachnus (see

pawlowae, Forda see hirsuta thunbergii), Pineus, Prociphilus (and
pawneepae, Aphis also see crataegicola), Stomaphis
paxilla, Cinara pinicorticis, Pineus see pinifoliae
pecanis, Monelliopsis pinidensiflorae, Cinara
pectinatae, Adelges, Cinara pinifoliae, Pineus
pelei, Betulaphis see brevipilosus piniformosana, Cinara
pendleburyi, Pseudoregma pinihabitans, Cinara,
pentandrinus, Chaitophorus pinikoraiensis, Cinara (see watanabei),
pepperi, Myzocallis Pineus cembrae ssp.
pergandei, Cinara, Prociphilus, piniphila, Cinara
Protopterocallis piniradiatae, Schizolachnus
perniciosa, Phylloxera piniradicis, Cinara
persicae, Brachycaudus, Lachnus piniradicivorus, Prociphilus
swirskii ssp., Myzus, Pterochloroides pinitabulaeformis, Cinara see
persicaecola, Brachycaudus see persicae formosana
persicina, Tetraneura see radicicola pinivora, Cinara
persimilis, Hemipodaphis, Paoliella pinona, Cinara
commiphorae ssp. piri, Nippolachnus, Sappaphis
petersoni, Cinara piricola, Schizaphis
petroselini, Dysaphis apiifolia ssp. pistaciae, Baizongia
phaenax, Eriosoma pisum, Acyrthosiphon
phaseoli, Trifidaphis see Smynthurodes pithecolobii, Anomalosiphum
betae piyananensis, Allothoracaphis
pheleodendroni, Chaitophorus plantaginea, Dysaphis
phenax, Pemphigus platani, Tinocallis
philippensis, Greenideoida platanoidis, Drepanosiphum
phlomicola, Brachycaudus see pilosus plicator, Rappardiella see Tuberaphis
photiniae, Sinomegoura takenouchii
phyllanthi, Amphorophora see Sitobion, plicatus, Pemphigus
Eonaphis, Macrosiphum see Sitobion plotnikovi, Brachyunguis
takahashii, Sitobion podocarpi, Neophyllaphis
physocaulis, Dysaphis podocarpini, Neophyllaphis
piceae, Adelges, Cinara, poketa, Cinara
Protohormaphis polychaeta, Myzocallis, Tetraneura
piceaerubensis, Prociphilus pomi, Aphis
piceana, Neomyzaphis see Elatobium ponderosae, Cinara
momii pontanilaricis, Adelges see laricis
piceicola, Cinara, Tsugaphis sorini ssp. popularia, Phylloxerina
pictus, Hoplocallis, Phylloxera populellus, Chaitophorus see populifolii
pilcornis, Cinara populeti, Chaitophorus
pilosula, Phylloxera populeum, Pterocomma
pilosus, Brachycaudus, Cinara, populi, Cornaphis, Doraphis,
Pachypappa, Pterocomma, Pachypappa, Pemphigus, Prociphilus
Sigmacallis see Pachypappa, see Epipemphigus niisimae,
Subtakecallis, Tuberculatus Pseudochermes see Phylloxerina
pinata, Cinara populialbae, Chaitophorus
pindrowi, Adelges populicaulis, Pemphigus
pinea, Cinara populicola, Chaitophorus
pineoides, Pineus populifoliae, Pterocomma
pineti, Schizolachnus populifolii, Chaitophorus
 Index to Species Names of Aphids 981

populiglobuli, Pemphigus pustula, Cavariella

populinigrae, Pemphigus pustulatus, Chaitophorus
populiramulorum, Pemphigus pyraria, Dysaphis, Melanaphis
populisucta, Ceratoglyphina see pyri, Aphanostigma, Dysaphis,
Doraphis populi Eutrichosiphum, Pyrolachnus,
populitransversus, Pemphigus Schizaphis, Sinomegoura
populivenae, Pemphigus pyricola, Eriosoma
populivorum, Neopterocomma pyrifoliae, Macrosiphum
prelli, Adelges
probosceus, Prociphilus quadrata, Protopterocallis
prolifera, Phylloxerina quadrimaculata, Brevicorynella
protospirae, Pemphigus quadrituberculata, Betulaphis
pruinosa, Cinara quaestionis, Allocotaphis
pruinosae, Chaitophorus, Lambersaphis querceti, Phylloxera and also see
pruni, Asiphonaphis, Eumyzus, Moritziella corticalis
Hyalopterus, Vesiculaphis querecus, Lachnochaitophorus,
prunicola (-us), Betacallis, Brachycaudus, Tuberculatus
prunicolus, Eumyzus querci, Stegophylla
prunifoltae, Ryoichitakahashia quercicola, Greenidea,
prunisuctus, Myzus see cerasi ssp. Hoplochaitophorus, Neothoracaphis,
umefoliae Stegophylla, Tuberculatus
pseudoalni, Myzocallis see Pterocallis quercifoliae, Eutrichosiphum,
pseudobyrsa, Asiphum see Pachypappa Greenidea, Stegophylla
pseudocaudata, Taiwanaphis quercifolii, Tuberculatus
pseudocoryli, Macrosiphum querciformosanus, Tuberculatus
pseudomontana, Pseudoregma see quercihabitus, Neosymydobius
koshuensis quercina, Phylloxera, Stegophylla see
pseudopasaniae, Eutrichosiphum see quercicola
pasaniae querciphaga, Betacallis,
pseudopopuleum, Pterocomma Eutrichosiphum, Greenidea,
pseudoschrankiana, Glyphina Heminipponaphis, Neothoracaphis,
pseudoschwartzii, Cinara Nipponaphis see Schizoneuraphis
pseudoshinae, Tiliaphis querciphila, Kurisakia
pseudotaxifoliae, Cinara quercus, Acanthochermes, Atarsaphis
pseudotsugae, Cinara see agrifoliae, Cervaphis,
pseudotsugii, Aiceona Diphyllaphis, Phylloxera, Stomaphis
psidii, Greenidea see formosana quinquemaculatus, Chaitophorus
pteleae, Mesocallis
pterothorax, Sinotherioaphis see radicicola, Stomaphis, Tetraneura
Chuansicallis chengtuensis radicivora, Cinara
puerca, Cinara radicola, Dysaphis
pulverulens, Cinara radisectuae, Tuberculatus japonicus ssp.
pumilae, Eulachnus ralumensis, Schoutedenia
pumilus, Chromocallis (see nirecola), ramani, Dysaphis
Mexicallis analiliae ssp. rameshi, Eutrichosiphum
punctata, Myzocallis ramicola, Chaitophorus
punctipennis, Euceraphis ramnifila, Aphis
punjabipyri, Schizaphis randii, Taiwanaphis
purpurea, Phylloxera symmetrica var. ranunculi, Dysaphis
pusillus, Chaitophorus, Neobetulaphis, rappardi, Neophyllaphis
Periphyllus see Trichaitophorus aceris rara, Cinara see pruinosa
982 Index to Species Names of Aphids

raychaudhurii, Eutrichosiphum sacharini, Drepanaphis

reaumuri, Dysaphis, Patchiella sacharinipini, Cinara
rebecae, Tuberculatus spiculatus ssp. sakurae, Tuberocephalus
recurvispinosus, Trichaitophorus salatigensis, Oregma see Astegopteryx
redelei, Phloeomyzus see passerinii pallida
remaudierei, Chaitophorus, saliapterus, Chaitophorus
Tuberculatus salicariae, Aphis
reticulata, Phylloxera salicicola, Cavariella, Chaitophorus (see
retipennis, Aiceona nigrae and matsumurai), Pemphigus
rhamni, Aulacorthum, Macchiatiella, (see saliciradicis), Phylloxerina,
Sitobion Pterocomma
rhamnicola, Aphis see mammulata salicifoliae, Elatobium see hidaensis
rhenanus, Periphyllus saliciniger, Chaitophorus
rhodendorfi, Pterocomma salicis ssp. saliciradicis, Fullawaya, Pemphigus
rhoicola, Aphis salicis, Aspidophorodon, Cavariella,
rhois, Carolinaia, Juncomyzus see Lachnus, Phylloxerina, Pterocomma
Carolinaia japonica, Melaphis salicorticis, Chaitophorus see
rhombifoliae, Pterocallis see alnifoliae macrostachyae
rhusae, Dasyaphis salicti, Aphis see Pterocomma
rhusicola, Kaburagia salicicola, Chaitophorus
rhusivermiciflua, Stomaphis salignus, Tuberolachnus
riccobonii, Forda salijaponicus, Chaitophorus, Clavigerus
rigidae, Cinara see Plocamaphis coreana,
rileyi, Eriosoma, Eulachnus, Pterocomma
Phylloxera saliniger, Chaitophorus
rimosalis, Phylloxera saltans, Tinocallis
ringdahli, Pterocomma see rufipes salviae, Sitobion
robiniae, Myzocallis see Appendiseta sambuci, Aphis
roboris, Lachnus sambucifoliae, Aphis see sambuci
robustiseta, Aiceona sanguiceps, Pterocomma
roepkei, Astegopteryx see basalis sankari, Eutrichosiphum
rogersoni, Dysaphis brancoi ssp. sanpunum, Pterocomma
rosea, Fushia see Nurudea yanoniella sanpupopuli, Epipemphigus
rosettei, Pachypappa sapporoensis, Tinocallis see
rostropunctata, Myzocallis takachihoensis
rotifera, Reticulaphis distylii ssp. saramaoensis, Neothoracaphis
rotunda, Metanipponaphis saraswatae, Cinara
roumanica, Aphis sarmaticum, Pterocomma pilosum ssp.
rubicunda, Cinara sasae, Takecallis
rufiabdominalis, Rhopalosiphum sasakii, Astegopteryx (see
rufipes, Pterocomma bambusifoliae), Prociphilus,
rufulum, Rhopalosiphum Tuberocephalus
rungsi, Geoica see utricularia sasicola, Yezaphis
ruperti, Hoplocallis saskensis, Cinara
russellae, Cinara, Eutrichosiphum, sassceri, Aphis
Phylloxera sauteri, Yamatocallis
rustica, Cinara sawagarumii, Kurisakia ailanthi ssp.
rutila, Phylloxera see coccinea sawashibae, Mesocallis
schimae, Greenidea
sabrinae, Drepanaphis schimitscheki, Cinara
sacculi, Pachypappa schinifoliae, Aphis
 Index to Species Names of Aphids 983

schlingeri, Neuquenaphis, Toxoptera similans, Phylloxera

and see aurantii similis, Calaphis, Capitophorus,
schneiden, Adeiges see nordmannianae Chaitophorus, Neuquenaphis,
schouteniae, Cervaphis Oregma see Astegopteryx bambusae,
schrankiana, Glyphina Pineus
schreiben, Myzocallis simlaensis, Eutrichosiphum
schuhi, Cinara simmondsi, Pineus see boerneri
schwartzi(i), Brachycaudus, Cinara simpsoni, Chaitophorus populifolii ssp.
scirpifolii, Rhopalosiphum see musae (see neglectus), Drepanaphis
sclerata, Tuberolachnus sinense, Eutrichosiphum
sclerophyllum, Eutrichosiphum see singaporensis, Astegopteryx see basalis
shiicola singeri, Periphyllus
sclerosa, Roepkea see Nearctaphis sinica, Nurudea see ibofushi
segregis, Adelges see viridis sininirecola, Chromocallis see nirecola
semenovi, Pemphigus see mongolicus sinipiricola, Sappaphis see pini
semiglabra, Nipponaphis sinipopulifoliae, Pterocomma see
semisubterraneus, Brachycaudus see yezoense
persicae siniquercus, Lachnus see tropicalis
sensoriatus, Chaitophorus populeti ssp., sinisalicis, Aleurodaphis,
Hyalomyzus, Nearctaphis, Aspidophorodon (see salicis),
Neuquenaphis Stomaphis
setacea, Doralis see Aphis lantanae sinobursarius, Pemphigus
setariae, Hysteroneura sinocaryae, Kurisakia
setigera, Astegopteryx, Parathoracaphis siphonella, Melanaphis
setosa, Cinara (see hyperophila), siphunculatus, Pemphigus
Glyphina sitchensis, Cinara
setulosa, Cinara, Geoica smaragdina, Cinara see todocola
shaposhnikovi, Chaitophorus, smithi, Muscaphis
Pachypappa smithiae, Pterocomma
sharmai, Dysaphis smolandiae, Cinara
shatungensis, Chaitophorus tremulae solanella, Aphis see fabae
ssp. solani, Aulacorthum
shepherdiae, Capitophorus solitaria, Cinara
shiiae, Lithocarpus, Neonipponaphis, sonata, Cinara
Reticulaphis sophorae, Tinocallis see insularis
shiicola, Eutrichosiphum, Lachnus, sophoricola, Aphis
Metanipponaphis sorbi, Dysaphis, Indotuberaphis
shinjii, Cinara, Mollitrichosiphum, sorbiarum, Dysaphis
Tiliaphis sorbiradicis, Eriosoma
shiraii, Nurudea sorini, Chaitophorus tremulae ssp.,
shirakabae, Hamamelistes see betulinus Cinara, Tetraneura, Tsugaphis
shirombana, Pachypappa see tremulae spicata, Drepanaphis
shitosanensis, Pseudoregma see spiculatus, Tuberculatus
koshuensis spiculosa, Cinara
siamensis, Aiceona see actinodaphnis spinifer, Mexicallis
sibirica, Dysaphis spiniferus, Hoplochaitophorus,
sibiricae, Cinara see juniperi Phylloxera see castaneae
sibiricus, Pineus see cembrae spinosus, Hamamelistes,
sichangensis, Forda Kaltenbachiella, Myzocallis,
sichuananus, Pineus Phylloxera see caryaecaulis
sikkimensis, Betacaliis, Eutrichosiphum spinuloida, Phylloxera
984 Index to Species Names of Aphids

spiraecola, Aphis Dermaphis japonensis, of Strand see

splendens, Cinara Dermaphis takahashii
spyrothecae, Pemphigus takaoensis, Paranipponaphis
stanfordiana, Phylloxera takenouchii, Astegopteryx see
stanleyi, Macrosiphum Tuberaphis, Tuberaphis
staphyleae, Rhopalosiphoninus tamariciarum, Brachyunguis see
steinheili, Pterocomma see rufipes tamaricis
stellata, Phylloxera tamaricifoliae, Brachyunguis see
steveni, Periphyllus tamaricis
stevensis, Chaitophorus tamaricis, Brachyunguis, Eulachnus see
stigmatus, Tuberculatus Eotrama
striata, Oregma see Astegopteryx tamaricophila, Brachyunguis
bambusae tamaricovorum, Brachyunguis see
strobi, Cinara, Pineus tamaricis
strobilobius, Adelges see laricis tanneri, Cinara
stroyani, Chaitophorus salijaponicus taoi, Eutrichosiphum
ssp., Cinara (see piceicola), tapatii, Eutrichosiphum
Muscaphis tarakoensis, Neothoracaphis
styraci, Hamiltonaphis tarani, Macchiatella rhamni ssp.
styracicola, Ceratoglyphina taraxacicola, Aphis
styracophila, Astegopteryx tardoides, Adelges see laricis
subapicula, Cinara tardus, Adelges see laricis
subellíptica, Phylloxera see caryaecaulis tasmaniae, Taiwanaphis
subinoyi, Eutrichosiphum tatakaensis, Lachnus
subnudum, Sitobion papillatum ssp. tattakana, Chaetoregma,
subterranea, Cinara Eutrichosiphum
sumatrana, Astegopteryx tauricus, Eulachnus see rileyi
sutepensis, Greenidea, Neothoracaphis taxifoliae, Cinara see pseudotsugae
suzhouensis, Tinocallis taxus, Prociphilus
swinhoei, Astegopteryx tegmentosus, Periphyllus
swirskii, Lachnus tenochca, Myzocallis
sylvestrii, Ceratovacuna, tenuicorpus, Mollitrichosiphum
Metanipponaphis tenuipes, Cinara
sylvestris, Pineus tepehuanensis, Myzocallis longirostris
symmetrica, Phylloxera ssp.
symplocois, Sinomegoura terminaliae, Paoliella
symplocosis, Greenidea terminalis, Cinara
syringae, Aulacorthum terricola, Fullawaya
szelegiewiczi, Chaitophorus tessellatus, Paraprociphilus
salijaponicus ssp., Shivaphis testacea, Aphis see frangulae
testudinatus, Periphyllus see
taedae, Cinara testudinaceus
taiwanus, Astegopteryx see Tuberaphis, texana, Phylloxera
Cinara, Mollitrichosiphum, thatcheri, Cinara
Takecallis, Tuberaphis theobaldi, Cavariella
takachihoensis, Tinocallis thunbergii, Eulachnus
takagii, Yamatocallis tianmushaensis, Tuberocephalus see
takahashii, Cavariella, Dermaphis, sasakii
Eutrichosiphum, Lachnus, Periphyllus tibetapini, Cinara
(see loricatus), Serratocallis, Sitobion, tibetasalicis, Pterocomma see rufipes
Stomaphis, Thoracaphis of Shinji see tibetensis, Pemphigus
 Index to Species Names of Aphids 985

tiliae, Eucallipterus, Macrosiphum, ulmosedens, Eriosoma see japonicum

Rhopalosiphoninus and also patchiae
tilicola, Eucallipterus see tiliae umbellulariae, Euthoracaphis
tilisucta, Shivaphis umefoliae, Myzus cerasi ssp.
tissoti, Drepanaphis, Hyalomyzus, unimaculata, Astegopteryx
Myzocallis, Schizoneurata uscare, Oregma see Pseudoregma
tistaensis, Cinara pendleburyi
titabarensis, Lachnus see Aiceona ushari, Brachyunguis see calotropicus,
todocola, Cinara ushikoroshi, Prociphilus
todomatsui, Adelges ussuricus, Prociphilus xylostei ssp.
tokyoensis, Periphyllus, Yamatocallis ussuriensis, Tinocallis see takachihoensis
tonaluca, Cinara utahensis, Chaitophorus, Drepanaphis
tonkawa, Myzocallis see Tuberculatus utricularia, Geoica
kiowanicus
torii, Adelges vagabunda, Cinara, Mordvilkoja
totarae, Neophyllaphis vagans, Anoecia
tragopogonis, Brachycaudus see vandenboschi, Macrosiphum,
schwartzi Periphyllus
trehernei, Pemphigus vandergooti, Metanipponaphis
tremulae, Chaitophorus, Doraphis vandermeermohri, Astegopteryx see
populi ssp., Gootiella, Pachypappa, Tuberaphis, Trichoregma see
Pterocomma Astegopteryx basalis, Tuberaphis
triangula, Tetraneura vandykei, Cinara
trochodendri, Elatobium variabilis, Boernerina, Cerataphis see
trojanae, Apulicallis fransseni
tropicalis, Lachnus varians, Myzus
truncatus, Chaitophorus variegatus, Chaitophorus
tschildarensis, Dysaphis varsoviensis, Pachypappa see
tsugae, Cinara warshavensis
tuberculatus, Callipterinella, vasculosa, Phylloxera see symmetrica
Indonipponaphis, Lizerius, venafuscus, Prociphilus see americanus
Monelliopsis, Myzocallis, venetiana, Periphyllus
Quernaphis, Tuberculatus see also venosus, Pemphigus
fangi verbasci, Aphis
tuberculifera, Phylloxera verhoeveni, Neopterocomma
tuberculostemmata, Eulachnus vermai, Clethrobius
tujafilina, Cinara versicolor, Aphis, Chaitophorus see
turajevi, Neopemphigus leucomelas
vesicalis, Pachypappa
ufuasi, Paoliella vesicarius, Pemphigus
uichancoi, Mesotrichosiphum victoria, Mindarus
ulmi, Eriosoma, Georgiaphis, victoriae, Toxoptera see odinae
Tetraneura villosa, Cinara
ulmicola, Colopha, Stomaphis, villosii, Periphyllus
Tinocallis viminalis, Chaitophorus
ulmifoliae, Aphanostigma see Olegia viminicola, Chaitophorus
ulmifolii, Tinocallis virgata, Dysaphis
ulmifusa, Kaltenbachiella virginiana, Patchia
ulmiparvifoliae, Tinocallis viridana, Adelges
ulmiphila, Mimeuria viridescens, Cinara see piceicola
ulmipumilae, Eriosoma viridipallida, Calaphis
986 Index to Species Names of Aphids

viridis, Adeiges, Betulaphis (see xylostei, Prociphilus

brevipilosa), Neophyllaphis,
Periphyllus, Tinocallis yamabiwae, Mollitrichosiphum
viridissima, Aphis yamatonis, Myzus
viridula, Adelges see viridana yanagicola, Aphis see farinosa
vitellinae, Chaitophorus yangcola, Pemphigus see borealis
viticis, Aphis yangi, Eriosoma
vittoriensis, Siculaphis yanoniella, Nurudea
voegtlini, Essigella see fusca yanonis, Neothoracaphis, Stomaphis
yezoense (-is), Pterocomma, Tetraneura
wahhaka, Cinara yinchuanensis, Tinocallis see saltans
wahlea, Schizolachnus see flocculosus yohoensis, Nearctaphis
wahluca, Cinara yokoyamai, Tuberculatus
wahsugae, Cinara see tsugae yomefuri, Chaitophorus populialbae
wahtolca, Cinara ssp.
wallichianae, Pineus yoshinomayai, Quadratus
walshii, Myzocallis yuccicola, Aphis see helianthi
wanepae, Cinara see vandykei yukona, Cinara see glehna
warshavensis, Pachypappa yulongshanense, Eutrichosiphum
watanabei, Cinara dubium ssp.
watsoni, Cinara yunglongensis, Lachnus
wertheimae, Geoica see utricularia, yunlanensis, Baizongia
Slavum yunnanensis, Callaphis nepalensis ssp.
westi, Cinara see Panaphis, Kurisakia, Pemphigus
wettsteini, Paoliella see Epipemphigus imaicus, Pterocallis
whiteshellensis, Aphis corylicola ssp.
wilhelminae, Illinoia
winforii, Patchia zachvatkini, Hoplochaetaphis
winonkae, Cinara see tujafilina zelkovae, Byrsocryptoides, Tinocallis
zelkovaecola, Byrsocryptoides
xanthii, Capitophorus zelkovisucta, Tetraneura see
xerophilae, Pterocomma Paracolopha morrisoni
xerophilaphica, Eotrama see Protrama zelkowae, Tinocallis
radicis zhangwuensis, Phloeomyzus passerinii
xitianmushanus, Nippolachnus ssp.
xizangensis, Chaitophorus (see inouyoi) zini, Dysaphis see anthrisci
Longistigma zoarcbursara, Cinara
 VII
PHOTOGRAPHS or SLIDE-MOUNTED
APHIDS

This series of photographs is the work of Mr P.V. York of the Photographic

Unit of The Natural History Museum.

987