Fauna of

New Zealand
Ko te Aitanga Pepeke
o Aotearoa

Klimaszewski, J.; Watt, J. C. 1997: Coleoptera: family-group review

and keys to identification. Fauna of New Zealand 37, 199 pp.
INVERTEBRATE SYSTEMATICS ADVISORY GROUP

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 Fauna of New Zealand
Ko te Aitanga Ρeρeke o Aotearoa

Number / Nama 37

Coleoptera:
family-group review
and keys to identification

J. Klimaszewski 1 and J.C. Watt 2

with illustrations by D.W. Helmore

Landcare Research
Mount Albert Research Centre
Private Bag 92170, Auckland, New Zealand

1Present address: BC Research Inc.

3650 Wesbrook Mall, Vancouver
British Columbia, Canada V6S 2L2
2
Research Associate

Manaaki
Whenua
PRESS
Lincoln, Canterbury, New Zealand
1997

1
Copyright © Landcare Research New Zealand Ltd 1997

No part of this work covered by copyright may be reproduced or copied in any form or bγ any means
(graphic, electronic, or mechanical, including photocopying, recording, taping information retrieval
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Cataloguing in publication
KLIMASZEWSKI, J. (Jan), 1950—
Coleoptera: family-group review and keys to identification /
J. Klimaszewski and J.C. Watt with illustrations by D.W. Helmore.
— Lincoln, Canterbury, N.Z. : Manaaki Whenua Press, 1997.
(Fauna of New Zealand, ISSN 0111-5383 ; no. 37).
ISBN 0-478-09312-8
I. Watt, J.C. Π. Title IlI. Series
UDC 595.76(931)

Prepared for publication by the series editor using computer-based text processing, Iayout, scanning, and
printing at Landcare Research, Mt AIbert Research Centre, Private Bag 92170, AuckIand, New ZeaIand

Māori text by Pacific International Translations Ltd, AuckIand

Published by Manaaki Whenua Press, Landcare Research, P.O. Box 40, LincoIn, Canterbury, N.Z.

Printed by GP Print Ltd, Wellington

Front cover: Huhu beetle, Prionoplus reticularis (Illustrator: D.W. Helmore).

PubIication of the Fauna of New Zealand series i s supported by the Foundation for Research, Science and
TechnoIogy under contract nos C09308 and C09617.
POPULAR SUMMARY HE WHAKAPOTONGA ΜΑ TE MAREA

Class Insecta
Order Coleoptera

Families of beetles

Beetles are the largest order of organisms, comprising over

350 000 described species in the world fauna. These spe-
cies are classified in some 23 000 genera. It has been esti-
mated that there are more species of beetles than there are Illustration /Whakaahua: Measurement and comparison
of vascular plants or fungi, and 90 times as many as there help us to classify beetles into families (Illustrator / Kai-
are mammal species. whakaahua: Kirsty Wilson).
In New Zealand at least 5182 native species of beetles
have been recognised. A further 343 species are either self- O ngā pūtoi raropi katoa, ko te pūtoi kurikuri te mea tino
introduced or have been deliberately brought in, for ex- nui. Neke atu i te 350 000 ngā momo kurikuri kua oti te
ample for the biological control of noxious weeds. These tautuhi, puta noa i te ao. E noho mai ana ēnei momo ki
beetles belong to some 1094 genera in 82 families. By ētahi puninga āhua 23 000 nei. E ai ki te whakaaro o
comparison the vascular plant flora includes about 2500 ētahi, he maha ake ngā momo kurikuri, tēnā i ngā momo
species, and the terrestrial vertebrate fauna about 350 spe- tipu whai iaia, i ngā momo kōpura whetū rānei, a, e 90 te
whakaraunga ake o ngā momo kurikuri i ngā momo kara-
cies. The actual number of beetle species in New Zealand
rehe whakangote.
is undoubtedly much higher — many are undescribed.
Neke atu i te 5182 ngā momo kurikuri o Aotearoa
The first European collections of New Zealand beetles
taketake ake kua kitea i tēnei whenua. Arā anō ētahi
were made by Joseph Banks and Daniel Solander during
momo e 343 nei tau noa mai, i āta kawea mai rānei, pērā
Captain James Cook's first voyage, in 1769, and the first
i ērā i whakanōhia mai hei pau i ngā taru kino. E noho
formal descriptions were by Johannes Fabricius in 1775. katoa mai ana ēnei momo ki ētahi puninga 1094, me ētahi
The origin of New Zealand's beetles is essentially un- whānau e 82. Kite whakaritea tēnei ki te nui o ētahi atu
known, but apparently at least some had developed on an rauropi, e 2500 ngā momo tipu whai iaia, ā, e 350 noa iho
ancient land mass of continental origin which became in- ngā momo kararehe noho whenua e whai tuarā ana. Kāore
creasingly isolated since the late Mesozoic era. The earli- e kore he nui noa am ngā momo kurikuri i Aotearoa i tērā
est known beetle fossil from New Zealand is of an incom- i huaina i runga ake nei, inā hoki, arā anō ētahi kāore anō
plete front wing dating from the Cretaceous. kia āta whakaahuahia, kia āta tautuhia.
Four factors are responsible for the composition and Ko Joseph Banks rāua ko Daniel Solander ngā Pākehā
diversity of the present-day beetle fauna of New Zealand. tuatahi i tahuri ki te āta kohikohi i ngā momo kurikuri o
(1) The origin of some of our fauna on the ancient south- Aotearoa, i a rāua ka haere mai i te rerenga tuatahi mai a
ern supercontinent known as Gondwana. (2) The estimated Hēmi Kuki ki Aotearoa i te tau 1769, ā, ko ngā tautuhinga
80 million years of geographic isolation of New Zealand, whaitake tuatahi, nil Johanns Fabricius i te tau 1775.
which has resulted in some 90 percent endemism (local Kāore e āta mōhiotia ana te pūtakenga mai o ngā
origin) of species, and high endemism also at genus level. kurikuri o Aotearoa, engari e ai ki te kōrero arā ētahi i
(3) Changing climate, land areas, and land surfaces due to takea mai i tētahi whenua rahi o tūāuriuri. Mai i te
factors such as glaciation, mountain building, and volcanic wāhanga whakamutunga o te wā Mesozoic, ka neke haere
activity. (4) The absence of mammals and other animals tēnei whenua rahi, mete aba, ka motuhake mai i ētahi atu
and plants which have dominated ecosystems elsewhere whenua. Ko te parawae kurikuri tino tawhito o Aotearoa
(continued overleaf) (ara haere tοnu)
in the world. The flora and fauna have been greatly modi- e mōhiotia ana, ko tētahi wāhanga o te pakihau o mua,
fied in the last 1000 years by humans, who brought devas- mai i te wā Cretaceous.
tation to native forest and introduced exotic animals with E whā ngā āhuatanga i whakatauhia ai te āhua me te nui
destructive influence (e.g., deer, rats, possum, goats). o ngā momo kurikuri o Aotearoa i ēnei ra. (1) Ko te
Contemporary New Zealand beetles are thus a compos- pūtakenga mai o ētahi momo i te whenua rahi tūāuriuri e
ite of ancient, variously changed lineages, elements intro- kīia nei ko Gondwana. (2) Ko te āhua 80 miriona tau e
duced by dispersal over short and long distances, and spe- noho motuhake mai ana a Aotearoa. Na konei i 90% ai te
cies intentionally and accidentally introduced by humans. nui o ngā kurikuri i Aotearoa nō konei taketake ake, ā me
The majority of species are associated with native forest, uaua ka kitea ēnei momo i ētahi atu whenua. Waihoki, kei
which now constitutes only 23 percent of the total land te pērā ano i te āhua ki ngā puninga. (3) Κo te rerekē
area. The shrinking forest has profoundly reduced the popu- haere o te āhua o ngā rangi, o te takoto mai o te whenua,
lation size of many native species and their potential for tae atu ki te mata o te whenua. Nā ngā āhuatanga pēnei i
long-term survival. ngā awa hukapapa, i ngā āhuatanga hanga maunga me ngā
Our beetles are also vulnerable to introduced organisms: mahi a Rūaumoko i pēnei ai. (4) Κo te kore o ētahi momo
their defence mechanisms are often ineffective against the karerehe whakangote me ētahi atu kararehe, tae atu ki
new arrivals. They may succumb to competition for food ētahi tipu kua noho mātāmua mai i ētahi atu rauwiringa
kaiao, hun i te ao. I tēnei 1000 tau kua hon ake nei, kua
with introduced wasps, to predation by mammals, and to
kaha tonu te rawekehia o ngā tipu me ngā kararehe o
depletion of food sources by weeds out-competing native
Aotearoa e te ringa tangata. Na te tangata kē hoki i kore
plants. A number of beetle species are recognised as being
hare ai ngā ngahere māori, nana anδ i tau mai ai te mahi
threatened with extinction, and are listed for protection by
a te kararehe kino i tawhiti (hei tauira, arā te tia, te kiore,
the Department of Conservation.
te paihamu, me te koti).
On a positive note, beetle diversity can be unexpectedly Na reira, ko ngā kurikuri o Aotearoa ināianei, i hua ake
high in relatively small areas of habitat. The beetle fauna
i te haere kōtui o ēnei āhuatanga e whai ake nei: ngā
of the Auckland suburb of Lynfield has been shown to kākano whakauru o tuauri whāioio, ngā mea i tau ā-
exceed 1000 species, of which three-quarters are native. puananī mai, ā-manapou mai rānei, ahakoa i ahu mai i pae
tawhiti, i pae tata rānei, tae atu ki ngā mea i āta haria mai
e te tangata, i haria pokerehū mai rānei e te tangata. Κo te
Contributor Jan Klimaszewski was born in Poland in nuinga o ngā momo nei, e honoa ana ki ngā ngāhere
1950. He graduated MSc in 1973 and PhD in 1978 from māori, ā, o te katoa o ngā whenua o Aotearoa, ko tētahi
the University of Wroclaw, with a specialisation in the 23% anake he wao māori. Na te korekore o ngā ngahere
beetle family Staphylinidae. He was based in Poland until māori, kua iti ake te maha o tēnā momo, o tēnā momo o
1980, when he took up a postdoctoral research fellowship ngā kurikuri tūturu o Aotearoa, ā, tērā pea kua kore e
in Ottawa, Canada. From 1982 to 1989 he was a research noho pūmau mai ki te mata whenua.
associate in the Lyman Entomological Museum at McGill Kua mōrearea anō hoki ā tātou ake kurikuri i te noho
University, working on lacewings (Neuroptera). Jan was tahi Id ngā rauropi kua tau mai i whenua kē, ia hoki kua
then appointed as a Senior Curator of Coleoptera in the kore e taea e rātou te kaupare atu ngā mate tērā pea ka pā
Transvaal Museum, Pretoria, South Africa. In 1993 he mai i ēnei rauropi hou. Tērā kua noho papa i te whaka-
returned briefly to Canada before taking up the post of taetae tahi ki ngā katip haere mai i tawhiti — e rua, e rua,
Coleopterist with Landcare Research in New Zealand. e whai ana i te kai kotahi; ko ngā kurikuri tonu rānei kua
noho hei kai ma tētahi kararehe whakangote; kua riro
Contributor J. Charles Watt was born in England but rānei ma ngā tarn tawhiti kē e tāmi ngā tipu o konei take-
moved to New Zealand at an early age. He graduated MSc take ake, me te aha anō, kua ēneipapāroa
tino kai a ngā
(Hons) in zoology from the University of Auckland in 1960. kurikuri.
In 1965 he graduated DPhil from Oxford University, where Arā ētahi momo kua tata tonu te korehāhā, ā, he mea
he worked on the beetle family Tenebrionidae. He then whakauru ēnei e Te Papa Atawhai ki te rārangi o ngā
joined the Entomology Division of DSIR, with responsi- rauropi kia āta maimoahia, kia āta rauhitia.
bility in the Systematics Section for all beetles except the Heoi anō, ia tirohia te taha rongo pai, arā ētahi wāhi
weevils. Charles was obliged to retire prematurely in 1986 noho ririki nei, ma kē te nui o ngā momo kurikuri e noho
because of a stroke which left him hemiplegic. He has since mai ana ki reira. E ai ki ngā kōrero, neke atu i te 1000 ngā
been an honorary research associate of the invertebrate momo kurikuri e noho tahi ana i te takiwā o Lynfield, i
systematics group, first under DSIR and since 1992 under Tāmaki-makau-rau, ā, ko tētahi 75% o ēnei, no konei
Landcare Research. taketake ake.
 PREFACE
The idea of producing a manual for identification of New Zealand beetle families originated long
ago in the Entomology Division of the former DSIR, yet for some time never quite got off the
ground. This changed when, after graduating in 1965 from Oxford University, Charles Watt joined
Entomology Division with responsibilities in the Systematics Section for Coleoptera. Over two
decades he gathered together representative specimens of many beetle species, compiling their
general descriptions, geographic distribution, and data on natural history, and produced a pre-
liminary linear key for the identification of our families. His selected examples, which have been
meticulously drawn by illustrator Des Helmore, represent most of the families occurring in New
Zealand. In October 1985, while working at the Natural History Museum in Paris, Charles suffered
a crippling stroke which culminated in his early retirement. This unfortunate event disrupted his
plans for completing the key project, although he has continued to work as a Research Associate
of Landcare Research.
In May 1994 Jan Klimaszewski arrived from Canada to work as a taxonomist on the beetle fauna
of New Zealand. His primary task was to check and update Charles Watt's key, but he opted to
prepare a manuscript reviewing New Zealand families, using many of Charles Watt's examples of
species. In consultation with Charles this was completed in late 1996, when the manuscript was
accepted for inclusion in the Fauna of New Zealand series. As first author, Jan was responsible for
preparing the manuscript, the present concept of the families, the family descriptions and their
arrangement according to the most recent system of classification, the data on nomenclature,
geographic distribution, and economic importance, the selected references, and line illustrations of
structures. He also produced a second, pyramidal key intended for quick identification. In late
1996 Jan took up an opportunity for employment in Canada, so as to be reunited with his wife
Krystyna and son Philip.
The published version has been considerably improved by the suggestions of several reviewers
and collaborators, all of whose input is warmly acknowledged, and with significant support and
guidance from the Fauna Management Committee (Marie-Claude Larivière, Trevor Crosby, Ross
Beever) and the Invertebrate Systematics Advisory Group (see p. ii). It has been structurally
developed and completed by the Series Editor, working from Jan's final draft, and integrating the
many suggestions of reviewers and collaborators.
This contribution attempts to fill a long-felt need, and critical feedback from users is invited, in
anticipation that a revised edition may incorporate many further improvements.

—Tymone Duval, Series Editor, June 1997

 WITHOUT TAXONOMY TO GIVE SHAPE TO THE BRICKS,

AND SYSTEMATICS TO TELL US HOW TO PUT THEM TOGETHER,

THE HOUSE OF BIOLOGICAL SCIENCE IS A MEANINGLESS JUMBLE.

— R.M. May, 1990: Taxonomy or destiny. Nature 347: Ι 29-130

DEDICATION
This work is respectfully dedicated to G. Kuschel, (1) J.F. Lawrence, ( w32)andAh.FoNest),Jr(
ground-breaking work on beetles of New Zealand and Australia, and on beetle classification in
general, encouraged this contribution and enabled its realisation.

(1)Dr Guillermo (`Willy') KuscheI, now retired, was a research scientist in CoIeoptera in the former Entomology
Division of DS1R, initially at Nelson and later at Mt Albert, AuckIand. He was largeIy responsible for the expansion
of the DSIR insect collection as a core element of the New Zealand Arthropod Collection. Willy is wideIy known
and respected for his research on the systematics of Curculionoidea.

(2)Dr John F. Lawrence is a Chief Research Scientist with CSIRO and Curator of Coleoptera at the Australian
National Insect Collection, Canberra. John is an outstanding coleopterist and a world leader in the higher
classification of beetIes. His work on Australian beetles, and his new system of classification, enabled us to
complete our present contribution.

(3)
Dr AIfred F. Newton, Jr is an Associate Curator and Head of the Division of Insects at the Field Museum, Chicago,
U.S.A., and a renowned specialist in the higher classification of Staphylinoidea. Al is interested in the world fauna
of beetles, and among many important papers has published on the New Zealand and Australian fauna.

— Jan Klimaszewski & Charles Watt

 ABSTRACT
The 82 beetle families represented in New Zealand are reviewed. These families are briefly
characterised in the phylogenetic order proposed by Lawrence & Newton (1995),1 with family
concepts .similar to those defined by these authors and by Pakaluk et al. (1994). 2 A general
introduction to the New Zealand beetle fauna is given, and collecting methods are briefly
discussed, and core literature is reviewed. Two alternative keys—pyramidal and linear—are
provided for identification of the families, using often different character sets. Morphological
features are explained, and many are illustrated. Estimated numbers of species, genera, and (where
appropriate) tribes and subfamilies are given. Known distributions are discussed, and data on
natural history and collecting are summarised. For each family at least one species is represented
by a habitus illustration, short description, and geographic profile, and important references are
listed. Keys are provided to subfamilies of Carabidae, Leiodidae, Staphylinidae, Trogossitidae,
Phloeostichidae, Scarabaeidae, Cerambycidae, Chrysomelidae, Anthribidae, Belidae, Brentidae,
and Curculionidae; for some other families a substantive publication containing a key is cited.
Some 1100 currently recognised genera are recorded for New Zealand, and approximately 5580
species (>5223 native, >3546 adventive) are recognised, including undescribed species housed in
the New Zealand Arthropod Collection. One endemic family, Chalcodryidae, and four endemic
subfamilies—Horelophinae (Hydrophilidae), Euderinae (Bostrichidae), Agapythinae (Phloeo-
stichidae), and Cyclaxirinae (Phalacridae)—are reported. An undescribed species of Platisus from
the Three Kings Islands is the only representative of the family Cucujidae in New Zealand.
1FamilesndubfoCptera(wihslcdgn,oterfcsadnmily-group
names). In: J. Pakaluk & A. Slipinski (eds), Biology, phylogeny and classification of Coleoptera: papers
celebrating the 80th birthday of Roy Crowson. Warszawa, Museum i Instytut Zoologii PAN. Pp. 779-1006.
2
Current classification and family-group names in Cucujoidea (Coleoptera). Genus 5(4): 223-268.

CHECKLIST OF FAMILIES
The checklist sequence follows mainly the classification Superfamily STAPHYLINOIDEA
proposed by Lawrence & Newton (1995). Numbers in [7] Family Hydraenidae 5 32 0 24
brackets are unique identifiers for each family. Columns of [8] Family Ptiliidae 13 >48 8/9 24
numbers are estimated totals of genera and species [9] Family Agyrtidae 1 2 0 25
currently recognised in New Zealand: col. 1 – genera; col. [10] Family Leiodidae 25 112 1 25
2– native species; col. 3 – adventive species. Numbers of [11] Family Scydmaenidae 11 >201 1 26
species are based on published data (e.g., Holloway 1982, [12] Family Staphylinidae 190 >936 >85 26
Watt l982a, 1992, Ordish 1984, Werner & Chandler (incl. Microsilphinae, Pselaphinae, Scaphidiinae)
1995, Klimaszewski et al. 1996) and the total number,
including undescribed species, in the New Zealand Arth- Series SCARABAEIFORMIA
ropod Collection (NZAC). Superfamily SCARABAEOIDEA
[13] Family Lucanidae 7 26 3 29
Suborder ADEPHAGA Page 1
[14] Family Trogidae 0 1 30
Superfamily CARABOIDEA [15] Family Scarabaeidae 25 132 12 30
[1] Family Rhysodidae 4 6 0 20
[2] Family Carabidae 75 >426 >19 21
(incl. Cicindelinae) Series ELATERIFORMIA
[3] Family Dytiscidae 11 16 0 22 Superfamily SCIRTOIDEA
[4] Family Gyrinidae 1 0 1 22 [16] Family Scirtidae 11 >125 0 31
(= Helodidae)
Suborder POLYPHAGA [17] Family Eucinetidae 1 1 0 32
Series STAPHYLINIFORMIA [18] Family Clambidae 2 9 2 32
Superfamily HYDROPHILOIDEA
[5] Family Hydrophilidae 25 70 5 23 Superfamily BUPRESTOIDEA
[6] Family Histeridae 13 23 6 23 [ 19] Family Buprestidae 2 2 1 33

-9-
 Superfamily BYRRHOIDEA [57] Family Corylophidae 8 >19 2 54
[20] Family Byrrhidae 8 >79 0 33 [58] Family Corticariidae 12 53 11 55
[21] Family Dryopidae 2 4 0 34 (formerly as Lathridiidae)
[22] Family Elmidae 1 16 0 34
[23] Family Limnichidae 2 7 1 35 Superfamily TENEBRIONOIDEA
[24] Family Heteroceridae 1 1 0 35 [59] Family Mycetophagidae 3 12 3 56
[25] Family Ptilodactylidae 2 6-8 0 35 [60] Family Archeocrypticidae 1 0 1 56
[26] Family Chelonariidae 1 1 0 36 [61] Family Ciidae 5 20 0 56
[62] Family Melandryidae 9 38 0 57
Superfamily ELATEROIDEA [63] Family Mordellidae 4 6 1 57
[27] Family Eucnemidae 7 22 0 36 [64] Family Rhipiphoridae 3 5 0 58
[28] Family Elateridae 26 >132 3 37 [65] Family Colydiidae 26 >196 0 58
[29] Family Lycidae 1 0 1 37 [66] Family Ulodidae 4 20 0 59
[30] Family Cantharidae 3 40 1 38 (formerly as Zopheridae)
[67] Family Chalcodryidae 3 5 0 59
Series BOSTRICHIFORMIA [68] Family Tenebrionidae 36 139 10 60
Superfamily DERODONTOIDEA [69] Family Prostomidae 1 1 0 61
[31] Family Derodontidae 1 2 0 39 [70] Family Oedemeridae 6 18 3 61
[71] Family Pyrochroidae 3 >7 0 62
Superfamily BOSTRICHOIDEA [72] Family Salpingidae 4 22 0 62
[32] Family Jacobsoniidae 2 3/4 0 39 (incl. Inopeplinae)
[33] Family Nosodendridae 1 2/3 0 40 [73] Family Anthicidae 7 17 9 63
[34] Family Dermestidae 7 11 6 40 [74] Family Aderidae 1/2 15 0 64
(formerly as Euglenidae)
[35] Family Bostrichidae 4 1 7 41 [75] Family Scraptiidae 2 >4 0 64
(incl. Lyctinae)
[36] Family Anobiidae 25 28 11 42 Superfamily CnRYSOMELOIDEA
(incl. Ptininae) [76] Family Cerambycidae 56 >180 8 65
[77] Family Chrysomelidae 33 >134 >19 66
Series CUCUJIFORMIA (incl. Bruchinae)
Superfamily CLEROIDEA
[37] Family Trogossitidae 10 >24 1 43 Superfamily CURCULIONOIDEA
[38] Family Chaetosomatidae 2 4 0 44 [78] Family Nemonychidae 1 4 0 67
[39] Family Cleridae 8 37 2 44 [79] Family Anthribidae 28 58 3 68
[40] Family Phycosecidae 1 1 0 45 [80] Family Belidae 3 11 0 69
[41] Family Melyridae 4 33 0 45 (incl. Aglycyderinae = Proterhininae)
(also as Dasytidae) [81] Family Brentidae 3 3 1 69
(incl. Apioninae)
Superfamily CUCUJOIDEA [82] Family Curculionidae 231 >1496 46 70
[42] Family Nitidulidae 12 21 11 46 (incl. Scolytinae)
[43] Family Monotomidae 2 1 5 46 Totals (approx.): 1100 >5223 >356
(incl. Rhizophaginae)
[44] Family Phloeostichidae 2 2 0 47
[45] Family Silvanidae 8 7 5 47 CONTENTS
[46] Family Cucujidae 1 1 0 48 Acknowledgments 11
[47] Family Laemophloeidae 2 3 3 49 Introduction 12
[48] Family Phalacridae 2 >3 1 49 Origins and affinities 12
[49] Family Cavognathidae 2 5 0 50 Composition of the N.Z. beetle fauna 13
[50] Family Cryptophagidae 12 18 12 50 Conservation status 13
[51] Family Languriidae 5 8 0 51 Morphology and biology 14
[52] Family Erotylidae 2 9 0 51 Study and identification of beetles 17
[53] Family Bothrideridae 2 6 1 52 Equipment and collecting methods 17
[54] Family Cerylonidae 2 >5 1 52 Accessing the literature: helpful references 18
[55] Family Endomychidae 3 6 1 53 How to use this review 19
(incl. Merophysinae) Review of families 20
[56] Family Coccinellidae 19 22 18 54 References 72

—10—
Glossary of technical terms 91 lionoidea and subfamilies of Curculionidae, and a list of
Keys to taxa: I `Pyramidal' key 97 genera for Curculionidae and Cerambycidae, and assisted
II `Linear' key 105 in gathering data on many families. Beverley Holloway
Illustrations: Fig. 1-100, schematic figures 113 contributed to sections on Lucanidae and Anthribidae.
Fig. 101-260, habitus figures 128 Rowan Emberson and John Marris (Lincoln University,
Examples of species (expanded captions) 168 Canterbury) contributed their knowledge to improve
Appendix 1: Threatened species of N.Z. beetles 184 various parts of the text, and provided many distribution
2: Beetle collections in N.Z. 184 records for the families. Rowan also assisted in correcting
3: Key to tribes of Carabidae 185 the difficult sections on Carabidae and Scarabaeidae. John
4: Distribution records for families 186 Lawrence (CSIRO, Canberra, Australia), Al Newton, Jr
Taxonomic index 188 (Field Museum, Chicago, U.S.), and Margaret Thayer
were extremely helpful in providing unpublished data,
sharing their knowledge, and commenting on the manu-
ACKNOWLEDGMENTS script. Laurent LeSage (Agriculture Canada, Ottawa,
Canada), Richard Leschen (Michigan State University,
This work is a compilation from previous taxonomic East Lansing, U.S.), Jyrki Muona (Finnish Museum of
revisions, occasional short papers, and beetle specimens Natural History, Helsinki, Finland), Masahiro Ohara
that have accumulated in the New Zealand Arthropod (Hokkaido University, Japan), Stewart Peck (Carleton
Collection over an extensive period. The specimens were University, Ottawa, Canada), and Clarke Scholtz (Uni-
collected primarily by G. Kuschel, J.C. Watt, J.S. Dug- versity of Pretoria, South Africa) provided help and
dale, B.A. Holloway, B.M. May, and many others not valuable suggestions. We express our appreciation to
mentioned but whose efforts deserve our sincere thanks. these collaborators, and to the many others who directly or
Des Helmore executed the great majority of the habitus indirectly contributed to our work.
drawings. His superb work is indispensable to us and to the For critical comments and helpful suggestions on
users of this publication; we salute him for his contri- various parts of the manuscript we thank Barbara Barratt
bution. The work of other contributing illustrators— (AgResearch, Invermay), John Bradshaw (Department of
Anthony Harris, Joanna Liddiard, Shaun Forgie, and Geology, University of Canterbury), George Gibbs
Lesley Alexander—is warmly acknowledged. We also (Victoria University, Wellington), Anthony Harris (Otago
appreciate the permission of John Lawrence and Jarmila Museum, Dunedin), John Hutcheson (Forest Research
Kukalová-Peck to reproduce figures first published in Institute, Rotorua), Chris Reid (James Cook University,
Lawrence & Britton (1994) and Kukalová-Peck & Queensland, Australia), and Greg Sherley (Department of
Lawrence (1993). Formal acknowledgment for illus- Conservation, Wellington).
trations is made in the captions (see especially p. 127). We are extremely grateful to the editor of the Fauna
Colleagues at Mt Albert Research Centre read the first series, Tymone Duval, for his enormous assistance in
draft and improved it with their comments: our thanks editing, formatting, and improving the final version of the
to Ross Beever, John Dugdale, Dianne Gleeson, and manuscript. His contribution was essential to achieve the
Research Associates Willy Kuschel and Brenda May. We present level of data organisation.
also thank Ross Beever for promoting an atmosphere of Jan Klimaszewski would like to thank his wife Krystyna
trust, creativity, optimism, and quiet scholarship. Marie- for her moral support and understanding during the pre-
Claude Larivière assisted with the computer literature paration of this work.
search on staphylinids, and provided the initiative and Charles Watt takes this opportunity to renew his
technical savoir fairetopermitn-huscagofe gratitude to Dr Roy A. Crowson (University of Glasgow)
habitus illustrations. Trevor Crosby helped in many ways for hospitality and intellectual stimulation. Many years
during the preparation of the manuscript; his influence has ago Dr Crowson visited New Zealand, and inspired
greatly improved the introductory sections and the Charles's interest in the family classification of our
references. Birgit Rhode checked and corrected German beetles. When Charles was studying at Oxford on the
references and their translations. André Larochelle helped relationships of tenebrionid beetles Dr Crowson invited
in organising major publications on our beetles according him to Glasgow, where he was able to study extensive
to family; he also assisted in gathering data on his collections of New Zealand tenebrionid larvae, and
specialty, the Carabidae, and checked and corrected the benefited from evening discussions on beetle family
French references and their translations. The friendly staff classification. Charles's manuscript linear key to the
of our Centre Library—Marleene Boyd, Jillian Irwin, and families of New Zealand beetles—circulated before he
Bella Smith—provided assistance with the references. went on study leave to Europe in 1984—was based on
We are most obliged to Willy Kuschel, who shared with Crowson's classification as it existed at that time.
us his extensive knowledge of New Zealand beetles, Charles wishes to especially acknowledge his father, the
provided keys to identification for families of Curcu- late Dr John S. Watt, who on learning of Charles's stroke
in 1985 flew to Paris and remained with him until he was instance the relict genera of tenebrionoid Pilipalpinae per-
able to be flown home. When Charles had recovered sisting in New Zealand, Australia, southern South Amer-
sufficiently, his father began to bring him twice weekly to ica, and Madagascar (Pollock 1995). The Chaetosomatidae
the Mt Albert Research Centre, where they would sit are known only from New Zealand and Madagascar, and
together preparing some of Charles's earlier work for the Phycosecidae from New Zealand, Australia, New
publication. This continued until John Watt, well into his Caledonia, and Vanuatu (J.F. Lawrence, in litt.). The small
eighties, was himself laid low by a stroke, and died in late family Cavognathidae, with species occurring in birds'
1996. The role of helper has been taken up by Charles's nests, is known from Australia, New Zealand, and Chile.
sister Mrs Heather Raudon. Charles is profoundly grateful The New Zealand Nemonychidae are closest to species
for this help, which was essential for him to be able to from Chile. Omaliine staphylinids of the genus Metacor-
complete his part of the present work from a wheelchair neolabium (22 species) are distributed in New Zealand,
and with impaired visual acuity. Australia, and South America (Thayer 1985). Some south-
This work was supported by the Foundation for ern temperate Staphylinoidea from families Hydraenidae,
Research, Science and Technology under contract nos Ptiliidae, Agyrtidae, Leiodidae, and Staphylinidae (esp-
C09308 and C09617. ecially Omaliinae and Pselaphinae) show `transaustral
disjunctions,' i.e., are isolated in two or more southern
land areas—New Zealand, Australia, South Africa, and/or
South America; for details see Newton (1985). In contrast,
INTRODUCTION the small, archaic family Agyrtidae, with fossil records
from the mid Jurassic (Newton 1991a), is distributed in the
Beetles are the largest order of organisms, comprising over
Holarctic region except for two species occurring in New
350 000 described species in the world fauna. They are
Zealand. Similarly, the staphylinid Stylogymnusa subant-
classified in some 23 000 genera (Endrödy-Younga 1985).
arctica is known only from the Auckland Islands, with its
Lawrence & Britton (1994) estimated that there are more
closest relatives exclusively Holarctic.
species of beetles than there are vascular plants or fungi,
The factors responsible for the composition and
and 90 times as many as mammal species. For a com-
diversity of the present-day New Zealand beetle fauna are:
prehensive definition of beetles see Lawrence & Britton
(1) the Gondwana origin of some of our biota; (2) the
(1994).
estimated 80 million years of geographic isolation of New
The first collections of New Zealand beetles were made
Zealand, which resulted in some 90% endemism at the
by Joseph Banks and Daniel Solander during Cook's first
specific level and a high level of generic endemism; (3)
voyage in 1769, and the first species were described by
changing climate, changing shorelines (Oligocene bottle-
Fabricius (1775). Watt (1979b, 1982a) and Ramsay &
neck), progenies, glaciation, and volcanic activity; and (4)
Singh (1982), amongst others, provide biographical notes
the absence of mammals and other animals and plants
about the early collectors and describers of beetles, and the
which have dominated ecosystems elsewhere in the world.
development of entomology in New Zealand.
The fauna has been greatly influenced in the last 1000
years by humans, who brought devastation to native forest
Origins and affinities
and introduced exotic animals with destructive influence
The exact origin of New Zealand beetles is essentially (e.g., deer, rats, possum, goat).
unknown, but there are indications that at least some Contemporary New Zealand beetles are a composite of
beetles had developed on an ancient landmass of con- ancient, variously changed lineages, elements introduced
tinental origin which became increasingly isolated since by dispersal over short and long distances (e.g., groups
the late Mesozoic (Daugherty et al. 1993). For example, dispersed passively over water as a result of the strong
the New Zealand jacobsoniid beetle Saphophagus min- westerly winds and currents around the 40th parallel), and
utus is considered by Crowson (1959) to be a relict coeval species intentionally and accidentally introduced by
with the tuatara (Sphenodon). humans. In support of the hypothesis that at least some of
Craw & Watt (1987) reported the first New Zealand our beetles are of ancient origin is the extraordinarily high
Cretaceous beetle fossil, an incomplete elytron, identified level of endemism at the specific level, with several
as belonging to a member of Polyphaga, either a buprestid endemic tribes, some endemic subfamilies, and many
or a chrysomelid. Kuschel (1987b) described subfossil endemic genera (Watt 1982a). Several families usually
fragments 1680 years old of a presumably extinct fern- small elsewhere are represented in New Zealand by
weevil, Tymbopiptus valeas, from near Te Kuiti, North diverse endemic forms (e.g., Byrrhidae, Colydiidae,
Island. For a recent review of the geological context of the Hydraenidae, Melandryidae, Ptiliidae, Scirtidae, and
evolution of the New Zealand biota, see Cooper & Scydmaenidae). The native element in our fauna is
Millener (1993). associated primarily with the lowland forest which has
In some groups present-day distribution patterns are prevailed throughout most of New Zealand's geological
suggestive of a Gondwana or pre-Gondwana origin, for history (Kuschel 1990), alpine habitats, tussock grass-

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lands, and the subantarctic islands. Introduced species are The most comprehensive survey ever conducted on
mostly associated with man-induced habitats (Kuschel New Zealand beetles is that of Kuschel (1990), in the
1990). suburb of Lynfield, Auckland, in which 982 species are
listed (the total number from the survey area, including
Composition of the New Zealand beetle fauna some collected after the survey was published, reaches
1000 — G. Kuschel, pers. comm.). Of these, 753 are en-
At least 5223 native and 356 adventive species of beetles demic and 229 adventive, and they belong to 65 families.
have been recognised in New Zealand, belonging to some Of the 753 indigenous species, approximately 10 are
1100 genera in 82 families (estimates based on currently restricted to beaches or intertidal wrack and 2 others to
recognised species housed in the New Zealand Arthropod beaches and open field situations. The great majority were
Collection and various museums: Watt 1976, 1982a, associated with native bush. Sixty-eight species, con-
Klimaszewski et a l.1996). The actual number of New stituting 9% of the 744 native non-littoral species in the
Zealand beetle species, including undescribed ones, is survey, were found in open environments but their sub-
probably much higher. By comparison the vascular plant sistence in these environments and their endemic status
flora includes about 2500 species, and the terrestrial were uncertain. Kuschel (1990) presented interesting
vertebrate fauna about 350 species (Watt 1976). Watt observations on the ecological flexibility of indigenous
(1976) estimated at 20 000 the number of described and (native) and foreign (introduced/ adventive) species. Of
undescribed species of terrestrial (including freshwater) the 982 Lynfield beetles, 55 belong to 16 genera with both
Arthropoda probably occurring in New Zealand. The native and foreign species. Of these 55 species, 22 natives
numbers are estimates, but nevertheless show that beetles were found only in the native bush, and all 33 foreign
constitute a significant proportion of New Zealand's total species occurred in home gardens and fields, but 14 of
insect fauna. these occurred also in the native bush environment.
The New Zealand beetles belong to two suborders, Kuschel (1990) believes that the beetle population of
Adephaga and Polyphaga. The suborders Archostemata Lynfield represents about 8-8.5% of the total (described
and Myxophaga, although present in Australia, do not and undescribed) New Zealand beetles—some 10 000 to
have representatives in our fauna. Of the 82 New Zealand 10 500 species. The conclusions of the Lynfield study
families, four—Archeocrypticidae, Gyrinidae, Lycidae, relate to all our beetles, which are predominantly forest-
and Trogidae—have only adventive species present. inhabiting species and show little interest in man-induced
Eleven families are represented by more than 100 des- environments.
cribed species. They are here listed in descending order, Ground-breaking investigations on the composition of
with approximate numbers in parentheses: Curculionidae beetle faunas in native forests and in degraded habitats,
(1542), Staphylinidae including former Pselaphidae and of beetles as indicators of habitat degradation, are
(1021), Carabidae (445), Scydmaenidae (202), Colydiidae being conducted by entomologists of the Forest Research
(196), Cerambycidae (188), Chrysomelidae (153), Scar- Institute in Rotorua (Hutcheson & Hosking 1994; Hutch-
abaeidae (144), Elateridae (135), Tenebrionidae (159), eson 1990, 1996; Hutcheson & Kimberley, submitted).
and Scirtidae (125). The numbers of Curculionidae,
Staphylinidae, and Carabidae in relation to the total num-
ber of beetle species in New Zealand are similar to the Conservation status
proportions in Australia and in the temperate Holarctic. The majority of New Zealand's beetle species are as-
However, the Colydiidae in New Zealand are particularly sociated with native forest, a habitat which has been
diversified, being better represented than elsewhere. Some drastically reduced by human activities over the past
smaller families, e.g., Byrrhidae, Cleridae, Hydraenidae, millennium. Native forest now constitutes only 23% of the
Corticariidae (=Lathridiidae), Leiodidae, Melandryidae, total land area. The `shrinking' forest has profoundly
and Trogossitidae, are also well represented in New Zea- reduced the population size of many native species and
land. There is one endemic New Zealand family, Chalco- their potential for long-term survival. Almost certainly
dryidae, and a few endemic subfamilies—Horelophinae many species have vanished before being documented,
(Hydrophilidae), Euderinae (Bostrichidae), Agapythinae and this process is probably continuing. Our beetles are
(Phloeostichidae), and Cyclaxyrinae (Phalacridae). In also vulnerable to introduced organisms because they have
comparison, there are approximately 20 000 described evolved in isolation, and their defence mechanisms are
beetle species in Australia (total beetle fauna estimated at often ineffective against the new arrivals (e.g., comp-
30 000) in 117 families; the largest are Curculionidae etition for food with introduced wasps, predation by
(6000), Scarabaeidae (3000), Chrysomelidae (3000), mammals, depletion of food sources by weed competition
Carabidae (2500), Staphylinidae including Pselaphinae with native plants).
(2500), Cerambycidae (1500), and Tenebrionidae (1200) The major threats to biodiversity are fragmentation and
(data from Lawrence & Britton 1994 and Klimaszewski et degradation of ecosystems due to inappropriate use of land
al. 1996). (e.g., deforestation) and water, invasion by exotic plants

—13—
and animals (Ramsay 1978), non-sustainable use of The capsule dorsally consists of vertex, frons, and cly-
species, and climatic changes (global warming, etc.). Con- peus (Fig. 1, 6); anterior and posterior tentorial pits, the
sequently populations associated with small, fragmented former (often called just tentorial pits) located at the
and degraded communities are particularly vulnerable, border of the clypeus and the frons; and ventrally the gula,
and are in need of conservation. The following citation genae, submentum, and mentum. The head is usually pro-
from Kuschel (1990) reflects the most important gnathous, i.e., with mouthparts facing forward (Fig. 1, 8),
conclusion from his study of the beetle fauna of the but may be hypognathous with mouthparts facing down-
Auckland suburb of Lynfield. ward (Fig. 9, 10). In many Curculionoidea the frons and
vertex are prolonged anteriorly to form a rostrum (Fig. 6,
Much of the native forest fauna is small, cryptic,
7).
elusive; and many species were rare for a start in
Compound eyes are usually well developed, except in
New Zealand, before the arrival of the Polynesians
some species living in permanent darkness (e.g., in deep
and, later, the Europeans. They are now on the way
soil or in caves). They may be coarsely or finely faceted.
to sure extinction unless the indigenous vegetation,
They are variable in size and shape (Fig. 8-12), and
which contains the special habitats that are so vital
sometimes (e.g., Gyrinidae, Tenebrionidae) each eye is
to their survival, is preserved. As this survey demon-
partially or completely divided into two by a lobe (Fig. 11)
strated, even the relatively small bush patches that so
called a canthus (Nichols 1989), ocular canthus, or genal
far have escaped destruction may contain faunas of
canthus (Watt 1988).
unsuspected richness and diversity.
The mouthparts (Fig. 4) are a labrum, two mandibles,
Examples of small and isolated beetle populations are two maxillae each comprising a lacinia, a galea, maxillary
an undescribed Platisus species known only from Great palps, glossae, and paraglossae, and a labium with labial
Island in the Three Kings group, Geodorcus ithaginis palps (Fig. 5). The glossae and paraglossae are usually
from Stack H in the Mokohinau Islands, and Stylogym- fused to form a median ligula. Variations in the terminal
nusa subantarctica, occurring exclusively on the Auck- segment of the maxillary palp (Fig. 13-16) have dia-
land Islands. The first is the only representative of the gnostic value for some groups.
family Cucujidae in New Zealand, and the last is the only The antennae are usually 11-segmented, with the basal
surviving gymnusine staphylinid (see p. 12) in the segment, or scape, articulated to the head and the second
Southern Hemisphere. segment, or pedicel, connecting the scape with the rest of
The Department of Conservation has provided a list of the antenna, called the flagellum (Fig. 1, 6). The last few
threatened species for New Zealand (Molloy et al. 1994) — segments may be swollen, forming a more or less compact
see Appendix 1. There are 26 native invertebrates, in- club. The main antennal types are:
cluding beetles, in category A (highest priority threatened filiform — the simplest form, with cylindrical segments
species), 52 in category B, and 20 in group C. The total which are nearly uniform in shape (Fig. 17, 18);
number of threatened invertebrates represent less than moniliform — with round, similar segments, resembling a
0.1% of the taxonomic groups in categories A—C (Molloy string of beads (Fig. 19);
et al. 1994). Examples of New Zealand beetles of in- serrate — with one side of each segment slightly pro-
determinate conservation status that are rare in collections truding, and forming a series of points like the teeth of
are Brounia thoracica (Chelonariidae), Lenax mirandus a saw (Fig. 20, 21);
(Monotomidae), and species of Microsilpha (Staphylinidae) pectinate — with one side of each segment protruding,
and Horelophus (Hydrophilidae). forming a series of long, sharp points like the teeth of
a comb (Fig. 22) (if both sides of each segment are
prolonged, the antenna is called bipectinate);
plumose — similar to the bipectinate antenna, but with the
MORPHOLOGY AND BIOLOGY projections on both sides of each segment slender and
flexible, like the vanes of a feather;
The basic morphological structures of beetles referred to clavate — with apical segment's enlarged into a club-like
in this contribution are shown in Fig. 1-3, which are expansion, either weak and gradual or abrupt and ball-
intended to be self-explanatory. We will, however, briefly like (Fig. 23-29);
characterise the major morphological features of the beetle capitate (a type of clavate antenna) — with apical segments
body. Readers are also recommended to consult the abruptly enlarged into a ball-like club (Fig. 23, 187);
Glossary (p. 91) for definitions of morphological terms. lamellate — with club segments having flat, leaf-like
projections on one side which usually can (Fig. 31) or
Head (Fig. 4-33). The head consists of a sclerotised cap- sometimes cannot (Fig. 30) be brought closely to-
sule, mouthparts, two compound eyes and sometimes one gether;
or two ocelli (e.g., Staphylinidae - Omaliinae, Derodon- flabellate — with long, thin lateral processes like alternate
tidae), and a pair of antennae. folds of a fan (Fig, 32, 33);

-14-
geniculate – with the second segment (pedicel) attached at Wings (Fig. 46-51). The first pair of wings are sclerotised
an angle to the first (scape), and each following seg- to form two elytra, which usually completely cover the
ment in line with the pedicel (Fig. 28, 140-143). abdomen (Fig. 1) or which may be reduced, exposing most
of the abdomen, as in the majority of Staphylinidae (e.g.,
Thorax (Fig. 1-3, 34-45, 52-56). The thorax consists of Fig. 132-138). The second pair of wings are the mem-
three parts—prothorax, mesothorax, and metathorax— branous flying wings (Fig. 46, 47), which are sometimes
each bearing a pair of legs lateroventrally. The mesothorax small or even absent, and are concealed by the elytra when
and metathorax are joined together to form the ptero- not in use. For details on venation and evolution of the
thorax, and each bears a pair of wings dorsolaterally. hind wings in Coleoptera (Fig. 48-51) see Kukalová-Peck
Each thoracic segment is made of four sclerites, with & Lawrence (1993).
their borders defined by sutures. The topmost (dorsal)
sclerite is the notum, the two lateral sclerites are the pleura, Legs (Fig. 1, 57-87). The beetle leg consists of a coxa
and the bottom one (ventral) is the sternum. Any one of the (usually with trochantin), a trochanter, a femur, a tibia,
thoracic sclerites is named by adding the appropriate and a tarsus. The coxa of each leg articulates with the
prefix: hence pronotum, propleuron/-pleura, prosternum, thorax at two points: the trochantin, which is the only
mesonotum, mesopleuron/-pleura, mesosternum, etc. The externally visible extension of the internal pleuron (Fig.
prominent part of the beetle thorax is the pronotum, which 57), and the pleuron, which is separated from the troch-
is variably shaped and is often referred to as the pronotal antin by a membrane. The trochanter is articulated to the
disc (Fig. 1). coxa (Fig. 58-66). Its junction with the femur may be
In Adephaga (also Archostemata and Myxophaga, not straight (Fig. 61) or slightly oblique (Fig. 60), but some-
represented in New Zealand) the pronotum is extended times is strongly oblique, as in Fig. 58 and 59 (e.g., in most
and deflexed ventrally on either side, forming the pronotal Tenebrionoidea, Bothrideridae), so that there is direct
epipleuron (Fig. 2, 34, 35). The pronotal epipleuron meets contact between the femur and coxa (heteromeroid
the lateral part of the pleuron (propleuron), which is trochanter). Each tarsus is divided into 1-5 segments
visible externally and forms part of the thoracic wall. The (tarsomeres or tarsites), and the last segment usually bears
pleuron (propleuron) is separated from the pronotal epi- two claws (Fig. 67-82), rarely one. The tarsal formula
pleuron by the notopleural suture, and from the sternum refers to the number of tarsal segments on each pair of legs.
by the pleurosternal suture. It may be divided into two
sclerites, an anterior proepisternum which forms part of Abdomen (Fig. 2, 3, 88-92). The abdomen consists
the wall, and a posterior proepimeron, which may be apparently of 10 segments in the male and 9 in the female,
extended to meet the sternum and closes (partially or each forming a complete or partial ring. The 9th segment is
completely) the coxal cavities from behind. The troch- modified, forming the genital segment, and in the male the
antin is a small sclerite separated (or not) from the pleuron 10th segment is usually much reduced or fused to segment
by a membrane and connected by the articulating condyle 9. Each abdominal segment is composed of four sclerites:
with the coxa (Fig. 57). a dorsal tergum or tergite, two pleura or pleurites (one on
In Polyphaga the deflexed lateral portion of the pro- either side), and a ventral sternum or sternite. A respiratory
notum is called the hypomeron, and is attached directly to spiracle opening onto each pleuron is the aperture of a
the lateral portion of the sternum because the pleuron has trachea. The spiracles are usually located in pleural mem-
become reduced, fused with the trochantin, and hidden brane, but in some Scarabaeoidea and Staphylinoidea they
internally (endopleuron) (Fig. 3, 36, 37). The hypomeron may be located on tergites.
is separated from the sternum by the notosternal suture; The visible part of the abdomen consists of five or more
the notopleural suture is absent as a consequence of sclerotised sternites (called ventrites) (Fig. 2, 3). The first
reduction and internalisation of the pleuron. The anterior sternite is absent, or represented by a small sclerite near the
portion of the sternum varies among Polyphaga. metacoxae; the second sclerite may be hidden behind the
The nota of the mesothorax and the metathorax are edge of the elytra, and is visible only in lateral view. In
divided into three sclerites: the scutum, scutellum, and referring to the first sternite we mean the first ventrite or
postnotum. The mesoscutellum, usually called just the the first visible sternite. In Adephaga the first ventrite is
scutellum, is visible at the base of the elytra (Fig. 1). The completely divided by the large and prominent metacoxae
ventral part of the pterothorax consists of two sterna and (Fig. 88, 89), but in the Polyphaga it is undivided and
the pleura (episterna and epimera) (Fig. 2, 3). The coxal continuous from side to side (Fίg. 90). The first ventrite
cavities are bounded anteriorly and posteriorly by the may bear a pair of straight or curved femoral lines behind
sternum and laterally by the pleural sclerites. The meso- or between the metacoxae (e.g., in Coccinellidae) (Fig. 92,
coxal cavities may be considered as laterally `open' when a). The abdominal ventrites are either movable and
partly enclosed by pleural elements (Fig. 42, 44) and separated from each other by an intersegmental membrane
laterally `closed' when closed by the sterna alone (Fig. 43, (Fig. 91), or some of them are connate, lacking an
45). intersegmental membrane but with the suture still visible

– 15 –
Table 1 Primary feeding mode of some beetle families Pupae are partly enclosed by the larval skin in surface-
and subfamilies (adults and larvae). feeding beetles, or pupation occurs within the food plant or
in cells in the soil. Some species make cocoons, which in
Predators certain exotic aleocharine staphylinids are constructed
Cantharidae, Carabidae, Cleridae, Coccinellidae, Coly- from silk (Ashe 1982).
diidae, Cucujidae, Lycidae, Staphylinidae, Trogossitidae Coleoptera larvae have a well developed and usually
Phytophages sclerotised head, three thoracic segments, and commonly
Apioninae, Chrysomelidae, Curculionidae, Nemonychidae, ten abdominal segments (rarely eight or nine). The first
Nitidulidae, Scarabaeidae thoracic segment is slightly enlarged and more heavily
sclerotised, abdominal segment 9 usually has paired
Scavengers or fungivores processes called urogomphi, and the 10th abdominal
Anobiidae, Anthribidae, Chalcodryidae, Ciidae, Corticari- segment is small and sometimes bears pygopods. Beetle
idae, Cryptophagidae, Dermestidae, Lucanidae, Ptiliidae, larvae usually have mouthparts adapted for biting and
Ptininae, Tenebrionidae chewing, as in the adults, and have usually from three to
six simple eyes, called ocelli or stemmata (Lawrence
1991). The larval body varies a lot but is usually cylin-
(e.g., Buprestidae). Occasionally the suture between the drical, and is either legless or with apparent appendages.
first two visible segments may be partially obliterated. The Many larvae which bore into plant tissue lack legs (e.g., in
last tergite of the abdomen when exposed forms the Curculionidae). Although beetle larvae are morphologically
pygidium. The reproductive organs are usually concealed very diverse, their mode of life has led to the development
within the tip of abdomen. of a few distinctive larval types; for details and illus-
The internal genital structures, although very important trations see Lawrence & Britton (1994).
for distinguishing species, are complex and are not dis- The main morphological categories of larvae are
cussed here; readers may consult Lawrence & Britton eruciform, scarabaeiform, apodous, and campodeiform
(1994). (Fig. 97-100). Eruciform larvae are cylindrical or some-
what flattened, with short legs and with either short uro-
gomphi or none. They are less active than campodeiform
Biology larvae. Many species of Chrysomelidae and Tenebrionidae
Adults usually are more conspicuous than larvae and occur have this type of larva. Scarabaeiform larvae are distinc-
virtually in all terrestrial habitats; they may be nocturnal or tive by their robust, C-shaped body and Iong legs. They
diurnal, and may or may not occur in the same habitat as occur in the soil or rotten wood, and are characteristic for
their larvae. Kuschel (1990) discussed habitat types and Scarabaeoidea. Apodous larvae have reduced antennae
the beetle fauna in the suburb of Lynfield, Auckland. The and palps and lack thoracic legs and urogomphi. This type
range of habitat niches even in a suburban environment is is typical for Curculionidae. Campodeiform larvae have a
overwhelming, and includes forest canopy, tree trunks, prognathous head, long and well developed legs, and com-
live wood, sound dead wood, decayed wood, wood-rotting monly long, unisegmented or multisegmented urogomphi
moulds, ground plants and fungi, leaf litter, compost, (Lawrence & Britton 1994). They are active and usually
dung, carrion, stream banks and beds, soil, beach sand, predatory. Most Carabidae and Staphylinidae have this
wrack in the intertidal zone, ponds, etc. type of larva.
Beetles feed on a variety of plant and animal material, Larvae are less conspicuous than adults and occur
and parts of living plants may be utilised (roots, stems, usually within their food source, e.g., inside timber
leaves, and flowers). Slime moulds, fungi, algae, decaying (Cerambycidae, Buprestidae, Anobiidae), inside stems
wood, leaf litter, carrion, dung, and other insects are food and leaves (Chrysomelidae, Curculionidae), in fruits
sources for various beetles. Feeding modes are summar- (Nitidulidae) and seeds (Bruchinae), in the soil and/or leaf
ised in Table 1. litter (Staphylinidae, Carabidae, Pselaphinae), in roots or
For information about the biology of the Coleoptera in dung (Scarabaeidae), in carrion (some Staphylinidae, e.g.,
general, see Crowson (1981). Aleochara), or on the ground near the base of plants.
Beetles are holometabolous insects, i.e., they undergo a Larvae of Coccinellidae occur on foliage, where they prey
complete metamorphosis. Adult beetles lay eggs which on soft-bodied insects. Many beetle larvae occur under the
hatch into larvae; frequently these occupy different bark of logs or in rotten wood (e.g., Silvanidae, Cucu-
habitats from the adults. When the larva has completed jidae), others are associated with fungi (e.g., Ciidae,
feeding and growing it enters a `resting' stage called the Cryptophagidae, Erotylidae), and some occur exclusively
pupa, in which the body of the larva is transformed into in birds' nests (Cavognathidae). Many larvae are of
that of an adult (metamorphosis). This enables particular economic importance because they compete for human
species to use different trophic niches (as larvae) and to food resources or damage human food products, and are
prevail in unfavourable conditions (as pupae). therefore regarded as pests, e.g., larvae of some Derm-

-16-
estidae, Elateridae, Scarabaeidae, Chrysomelidae, Curculio- connecting it with the lower hoop. The latter is attached to
nidae, Anobiidae, Cerambycidae (see Booth et al. 1990). a sleeve 1 metre long which can be tightened at the bottom.
A family-level treatment of the larvae of New Zealand Sifters are useful for collecting beetles from all kinds of
beetles by Dr J.F. Lawrence (CSIRO, Australia), with keys ground litter, rotting wood, fungi, shore debris, lichens,
for their identification, is anticipated to be published in our mosses, leaf mould, and birds' nests. The beetles may be
Fauna series. Consequently mention of larvae in this work picked from sifted material placed on white sheets or large
has been kept to a minimum. white trays, or the sifted debris may be placed into a
Berlese funnel or other separator.

Berlese funnels. The Berlese funnel in its various modi-

STUDY AND IDENTIFICATION OF BEETLES fications is used to separate insects from sifted debris. The
Important repositories of New Zealand beetle collections sifted sample with insects is placed on a screen near the top
are identified in Appendix 2, with contact details. of a funnel which is closed by a cover. An incandescent
bulb above the sample produces light and heat, drying the
litter and driving the insects downwards into the funnel,
Equipment and collecting methods
which ends with a container at the bottom. The container
Beetles occur in virtually all terrestrial habitats, therefore may be empty for catching live insects, or may contain a
collecting them requires knowledge of many different killing agent.
techniques. The most common and effective methods for
collecting beetles in various environments are listed Flight interception traps. The basic principle of such
below. For more details on collecting and mounting traps is to catch flying beetles by placing a screen in their
specimens see Martin (1977), Steyskal et al. (1986), way. The most successful interception traps tested in New
Walker & Crosby (1988), and Kuschel (1990). Zealand are the window trap, consisting of a vertical net
with a trough containing ethylene glycol underneath, and
Hand-collecting. Mainly medium-sized to large specimens the Malaise trap, which consists of a vertical net serving as
may be collected by hand from a variety of habitats (e.g., a baffle, end nets, and a sloping canopy leading up to a
from under stones, bark, rotting wood, foliage, etc.). collecting container. Attractants may be used to increase
the efficiency of these traps, of which there are many
Collecting nets. There are three basic types of nets: aerial varieties.
nets for collecting flying beetles, sweeping nets for
sweeping vegetation, and aquatic nets for gathering Pitfall traps. Pitfall traps are very successful devices for
insects from water. Aerial nets are light and the most collecting ground-walking beetles, or flying beetles if a
fragile. Sweeping nets are more robust, made of strong bait is used. They consist of an open-topped container
material with reinforcement around the metal ring to sunk in the earth. A cover may be placed over the open top
withstand being dragged through dense vegetation. Water to prevent rain flooding, but open space is required
(aquatic) nets are made of metal screening or heavy scrim between the cover and the rim of the container. The trap
with a canvas bag affixed to a metal rim. may be one-third filled with ethylene glycol preservative,
and the bait (carrion, dung, or a specific insect attractant
Beating sheets. A beating sheet is made of strong cloth e.g., molasses, beer, fruits, cheese) may be suspended
attached to a frame about 1 metre square, with two pieces inside the trap above the surface of the glycol. Trapped
of light wood or metal crossing each other and fitted into a insects should be collected regularly from the glycol
pocket at each corner of the cloth. The beating sheet is held solution and placed in 75% ethyl alcohol/acetic acid
or secured under a shrub or tree, and the branches or mixture.
foliage are beaten with a stick. Specimens will fall onto the
sheet, and can be removed using fingers, forceps, or an Light traps. An effective way to collect nocturnal flying
aspirator. (An inverted umbrella can also serve as a beating beetles is to use artificial light as an attractant and capture
sheet.) the beetles around the light. A mercury-vapour lamp,
fluorescent black light, or other lamps high in ultraviolet
Sifters. Sifters are containers made of cloth, wood, or wavelengths usually give excellent results. The light
plastic with a wire-mesh bottom serving as the sifter. The source may be placed against a vertically hung white sheet,
size of mesh used depends on the size of the specimens with a bottom sheet spread on the ground underneath for
sought. The most common model of sifter used in Europe falling beetles. More sophisticated light traps consist of a
and North America is made of two hoops of metal about 30 funnel with a killing container, and a suspended light with
cm in diameter, each provided with a handle. The lower vertical rigid screen most often made of transparent
hoop bears a wire-mesh screen welded to its edge. The top plastic. The flying insects approach the light, hit the
hoop is attached to a canvas cylinder about 30 cm long screen, and are collected in the killing container.

17—
—17---
Aspirator. An aspirator is a simple suction apparatus for publications form a core of early literature, and central to
picking up insects. The most common model consists of a them are the seven volumes of his Manual of New Zealand
vial of glass or transparent plastic with a close-fitting Coleoptera, published in four parts (Broun 1880, 1881,
rubber stopper. Two rubber or plastic tubes pass through 1886, 1893b). His work is listed here in extenso, even
the stopper. One is attached to a rubber tube through which though it may be of only limited value to workers today.
suction is created (by mouth, or with a rubber bulb) while • Chapman (1971): textbook on morphology, physiology,
the other end is filtered. The second tube extends approx- and natural history of insects.
imately 10 centimetres from the stopper, and sucks the
insects in. This device acts as a miniature vacuum cleaner, • Cooper (1979): review of fossil Coleoptera.
and is very useful for collecting small beetles. • Crosby & Larochelle (1994): preliminary list of Coleo-
ptera genera and families represented in the New Zealand
Fogging. Fogging with insecticides is the most recently Arthropod Collection.
developed method for collecting beetles from the tree
canopy, branches, trunks, or moss-covered surfaces. For • Crowson (1955): major treatise on the natural classi-
details, see Erwin (1983), Allison et al. (1993), and fication of the families of Coleoptera, based on detailed
Lawrence & Britton (1994). morphology of adults and larvae. Purported to reflect the
phylogeny of the Order, and gained wide acceptance.
Killing and relaxing. Beetles should be killed in jars with Includes keys to superfamilies, families, and some sub-
a little ethyl acetate absorbed into a suitable material (e.g., families. A cornerstone in the classification of beetles,
coarse hardwood sawdust such as oak, or crumpled variously modified in Crowson's later papers (1960, 1967,
absorbent paper) inside the killing jar. The absorbent l97la, b, 1973b). See Lawrence & Britton (1974) for
material acts as a barrier between the insect and the killing literature.
agent. An alternative solution is 75% alcohol with a little • Crowson (1981): textbook on biology of Coleoptera.
acetic acid or commercial vinegar to prevent stiffening of
• Hudson (1934): the first and only representative cata-
the insect's muscles. Ventral structures obscured by in-
logue of New Zealand beetles and their larvae, with many
conveniently placed legs in set specimens can be exposed
colour plates. Forms a watershed between the earlier
by soaking the specimen for a few minutes in Barber's
pioneering work and subsequent studies. An essential
fluid, or water with ammonia, to soften the joints suffici-
reference point.
ently for appendages to be moved without damage.
• Kuschel (1990): exhaustive study of the beetle fauna of
Accessing the literature: helpful references an Auckland suburb, remarkable for identifying almost
The literature on New Zealand beetles is extensive but 1000 species from native bush and modified habitats.
largely fragmentary. It is therefore necessary to know how Many habitus illustrations by Des Helmore, especially of
to access relevant publications. During compilation of weevils; also tabular summaries of species data, inform-
references for this review it became apparent that a few of ation on collecting methods, habitat types. A primary
them are of outstanding general relevance. Our selection is source of data included in the present work, and an
offered here in alphabetical order, since any attempt to essential resource for the student of New Zealand's
categorise in order of perceived value would be un- Coleoptera. *
realistic. However, it is fair to say that a comprehensive • Lawrence (1982): synoptic review of beetle families
recent bibliography, a general textbook or two on beetle with family limits and changes to classification.
systematics and biology, and regional summaries both
• Lawrence (1991): discussion, key, and many family
general and faunistic must be almost indispensable. Any
treatments of the immature stages of Coleoptera.
electronic reference sources—CD-ROMs, Internet web-
sites, searchable databases—must be seen as a bonus at the • Lawrence & Britton (1991, 1994): the Coleoptera chap-
time of writing, though these will undoubtedly grow in ter in the primary reference book Insects ofAustralia, and
prominence. its spin-off Australian Beetles, both with much relevance
The following publications, then, are so relevant to the to New Zealand.
present context that we have not repeated them under the • Lawrence & Newton (1982): review of the evolution
family treatments. They should be borne in mind as a first and classification of beetles at the superfamily level.
point of reference. Discusses origins and relationships, early fossils, sub-
• Böving & Craighead (1931): the first comprehensive ordinal classification, and evolution of Adephaga and
textbook on morphology of larvae. Polyphaga.
• Broun (1876-1923): Thomas Broun's name is appen- • Lawrence & Newton (1995): comprehensive current
ded to more species of New Zealand beetle (and, regret- classification of Recent Coleoptera at subfamily level and
tably, synonyms) than any other author's by far. His above, with complete or selective list of genera included in

—18—
each group. Discusses recent changes to classification, We also draw attention to the following publications
with extensive reference to studies providing evidence in reviewing beetles from other zoogeographical regions.
support of these changes. • AUSTRALIA: Matthews (1980-92), Lawrence & Brit-
• Meads (1990): introduction to rare, endangered, and ton (1994), Semmens et al. (1992).
protected invertebrates in New Zealand. • NEW GUINEA: Gressitt & Hornabrook (1977),
• Neave (1939-50): the Nomenclator Zoologicus with its Gressitt & Szent-Ivany (1968).
supplements is the definitive catalogue of genus-group • PACIFIC ISLANDS: van Dyke (1953), Nishida (1994),
names in zoology. Peck & Κukalová-Peck (1990).
• New (1995): textbook on conservation biology of in- • NORTH AMERICA: Bousquet (1991a), Arnett (1973),
vertebrates; the first to give a global overview of all major White (1983).
habitats (terrestrial, freshwater, marine). • SOUTH AMERICA: Solier (1849), Sharp (1882-87),
• Nichols (comp.) (1989): a revised edition of the well Blackwelder (1957).
known and comprehensive Torre-Bueno glossary of ento- • SOUTHERN AFRICA: Scholtz & Holm (1985).
mological terminology. • GENERAL: Evans & Bellamy (1996).
• Pakaluk & Slipinski (eds) (1995): a collection of the Important references on beetle larvae are Arndt (1993),
most recent papers on biology, physiology, and classi- Booth et al. (1990) (pest species), Böving & Craighead
fication of Coleoptera. (1931), Hudson (1934), Lawrence (1991) (key to fam-
• Ramsay & Crosby (1992): bibliography of New Zealand ilies), Lawrence & Britton (1991, 1994), Lawrence &
entomology up to 1985, and a guide to the bibliographic Newton (1982, 1995), Lawrence et al. (1993) (beetle
database BUGS, which is searchable on a `by family' basis larvae keyed on CD-ROM), May (1966, 1993) (larvae of
[accessible on application to the Curator, NZAC, email Curculionoidae), Stehr (1991).
address crosbyt@landcare.cri.nz ]; see also Crosby &
Ramsay (1992, 1994). How to use this review
• Ramsay & Singh (1982): concise guide to entomology The section Review of Families is presented in systematic
in New Zealand, including an historical review. sequence—notionally reflecting a natural phylogeny—of
• Shapiro & Porter (1989): paper reviewing the signi- numbered families, as in the Checklist of Taxa. Groupings
ficance of genitalia in the interpretation of insect evolution from suborder to superfamily are briefly introduced as a
and systematics. framework to this systematic treatment.
Under each family [numbered in brackets, for ease of
• Snodgrass (1935): textbook on insect morphology, use- cross-reference throughout] any assistance received from
ful for naming of external and internal structures. specialist col aborators is acknowledged. The next line
• Walker & Crosby (1988): concise manual on the identifies the habitus figure(s) representing that family.
preparation and curation of insect specimens.* Other relevant illustrations are not listed here, but appear
in the text wherever appropriate. Next, the range of body
• Watt (1956-92): Charles Watt's published work is listed length—measured from the apical margin of the labrum to
here in full, in view of its great relevance to the modern the apex of the abdomen—for members of the family is
study of Coleoptera in New Zealand. His Presidential given, along with the tarsal for-mula. This information
Address to the 1981 conference of the Entomological alone can be helpful in delimiting certain families or
Society of N.Z., New Zealand beetles (Watt l982a), is a groups of families.
valuable review of the field and a primary source of Subsequent information for each family is listed under a
information for the present work. sequence of subheads. The well defined families are ac-
• Wilson (1971): review of social insects and their affili- companied by a relatively short Diagnosis and Synopsis,
ations with other insects, including beetles. but those represented by more diverse forms have required
• Winterbourn & Gregson (1989): guide to the aquatic longer treatments. Character states in bold type constitute
insects of New Zealand, including the five or six beetle the primary diagnosis; the balance are supplementary. It
families with aquatic or hydrophilous habits as adults and/ will be noted under Synopsis that the family is referred to
or larvae. by the contracted form of its scientific name, as for ex-
ample Carabidae / carabid(s). This informal usage may be
preferred to colloquial names suggested under Remarks.
Range is represented as `recorded' (signified by ®) or
*AvaiIable from Manaaki Whenua Press, P.O. Box 40, Lincoln `unrecorded' (O) in the three major and eight minor
8152, New Zealand, tel. (03) 325 6700, fax (03) 325 2127, email islands or island groups constituting New Zealand; it is
mwpress @ landcare.cri.nz . shown for all families alphabetically in Appendix 4.

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 Under Examples, species representative of the family or REVIEW OF FAMILIES
constituent subfamilies are listed. Authorship and year are
cited, so that original descriptions may be pursued. Figure Suborder ADEPHAGA
numbers give access to both the habitus illustration and its We have followed the concept of Lawrence & Britton
expanded caption (under Examples of Species), which (1994) in defining this suborder: 1st abdominal sternite
offers a profile of the species illustrated. divided by metacoxae (Fig. 2); abdominal sternites 6 in
Under Remarks are given any additional points of number; pygidial defence glands present; larvae char-
information thought to be helpful, including reference to acteristic (for details see Lawrence & Britton 1994). The
any revisions known to be in progress, followed by a great majority of species are predacious. The phylogeny
suggested colloquial name. Some of these names are well and evolutionary history of the Adephaga is discussed by
established internationally, others are acceptable options Beutel (1995). The Adephaga comprise four families in a
in New Zealand. single superfamily, all represented in New Zealand.
The major families are furnished with a Key to their
subfamilies, or with references to such keys.
Finally, under Selected References we offer a list of Superfamily CARABOIDEA
publications considered likely to be most helpful in further This superfamily consists of families Rhysodidae, Cara-
exploring the family under consideration. These have been bidae (including Cicindelinae), Dytiscidae, and Gyrinidae.
kept to a minimum in order not to deter the more casual The majority of species, except for mycophagous Rhy-
user. Fuller bibliographies can be found elsewhere, as sodidae, are predacious. Rhysodidae and Carabidae are
indicated above, although for some families very little has terrestrial, Dytiscidae and Gyrinidae are aquatic.
been published that might be helpful in a New Zealand
context.
Morphological structures named in the text are intro- [1] Family RHYSODIDAE
duced in the section Morphology and Biology (p. 14); Fig. 101
most are also illustrated. A Glossary (p. 91) defines most
Length 5-10 mm Tarsal formula 5-5- 5
morphological terms, along with many others used in this
publication that may be unfamiliar. DIAGNOSIS. Body narrowly elongate, subcylindrical
Two alternative dichotomous keys are provided for to slightly flattened, widest at middle of elytra, glabrous
identification of New Zealand Coleoptera. They represent (except for tactile setae), black or exceptionally reddish
slightly different concepts in identification, and are meant brown, strongly glossy. Head grooved and strongly
to enable users to confirm an identification by using both. constricted posteriorly to form a narrow neck. Ant-
The pyramidal Key I (p. 97) is divided into several ennae stout and beadlike (Fig. 19). Pronotum with 1-3
components, and groups families sharing distinct features deep, longitudinal grooves. Legs short, robust; procoxae
regardless of their systematic relationships; it represents globose, their cavities closed.
the shorter identification route, and is intended for rapid or
preliminary identification. The linear Key II (p. 106) is not SYNOPSIS. There are 6 species of Rhysodidae known
subdivided, and represents slightly the longer identification from New Zealand, in 4 genera, of which 2 are endemic..
route. The two keys often employ different character sets. Adults and larvae are thought to feed on plasmodia of
They are designed to identify the families of the New Myxomycetes (J.F. Lawrence, pers. comm.), and live in-
Zealand coleopteran fauna, and may not be satisfactory for side dead or rotten wood. They may be found under bark or
other faunas. Characters which require dissection or dis- in wood of logs, standing dead trunks, or stumps in a partly
memberment of specimens have been avoided. Both keys decayed but not excessively wet or dry condition.
have been thoroughly tested, but may still contain errors or Collecting: sifting leaf litter, hand picking from under
inconsistencies. In the pyramidal key some families with bark, or crumbling and sifting pieces of moderately rotten
characters which might be overlooked, or are dubious, or old logs.
which differ between the sexes reappear under different
Groups. Identifications derived from the keys may be RANGE. North ®, South ®?, Stewart O
checked against the appropriate habitus illustration(s) and, Kermadecs O, Three Kings O, Chathams O, Snares O
if des-ired, the family diagnosis. Cross-reference to figure Aucklands O, Campbell O, Antipodes O, Bounties O
num-bers and family numbers is provided for convenience.
For the benefit of anyone using this publication exten- EXAMPLE. Rhyzodiastes proprius (Broun, 1880) (Fig.
sively for identifications at the microscope, we strongly 101).
recommend that you photocopy either the keys or the
illustrations (or both), and use them conjointly flat on the REMARKS. The Rhysodidae are sometimes treated as
bench-top. Single copies for this purpose are not in breach tribe Rhysodini of Carabidae.
of the publisher's copyright. Suggested colloquial name: heraldic beetles.

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 SELECTED ®,
®,REFERENCES. Bell & Bounties
Bell (1978, 1979,
Antipodes Aucklands ®, Campbell
1982, 1985), Bell (1991), Beutel (1990), Emberson
(1995), Watt (1980a). EXAMPLES. CICINDELINAE - Neocicindela tuberculata
(Fabricius, 1775) (Fig. 102); CARABINAE - Maoripamborus
fairburni Brookes, 1943 (Fig. 103); MIGADOPINAE - Lox-
[2] Family CARABIDAE omerus nebrioides Guérin -Méneville, 1841 (Fig. 104);
(incl. CICINDELINAE) SCARITINAE - Clivina basalis Chaudoir, 1843 (Fig. 105);

[assisted by R.M. Emberson and J.F. Lawrence] ΤRΕCΗΙΝΑΕ - Zecillenus alacris (Broun, 1821) (Fig. 106);
HARPALINAE - Ctenognathus novaezelandiae Fairmaire,
Fig. 102-107 1843 (Fig. 107).

Length 1.4-39 mm Tarsal formula 5-5-5
REMARKS. The most recent subfamily classifications of
DIAGNOSIS (see Fig. 1, 2). Body generally more or Carabidae have been proposed by Erwin (1991), Beutel
less streamlined, with series of punctures of fixed (1992, 1993), Arndt (1993), and Lawrence & Newton
position bearing tactile setae, usually glossy and black (1995). In older classifications the subfamily Cicindelinae
but in some species colourful and spotted. Head usually was regarded as a distinct family.
flattened, prognathous, with sharp, strong mandibles Bousquet & Larochelle (1993) proposed a classification
facing forwards. Antennae 11-segmented, usually fili- of Carabidae with 17 tribal divisions only, because of the
form, inserted between eyes and base of mandibles lack of consensus among authors at the subfamily and
(except Cicindelinae). Eyes prominent. Thorax smooth, supertribe level. Dr J.F. Lawrence has provided a key to
with lateral margins often well defined. Elytra simple, tribes of Carabidae in New Zealand (Appendix 3); This
usually sculptured with longitudinal grooves (striae). may be preferred by some users over the subfamily key
Legs usually strong, suited for running; protibiae with below.
apical antennal cleaning organ consisting of an exca- No comprehensive treatment of New Zealand Cara-
vation lined with a comb-like setal fringe; procoxae often bidae is available. However, the group is under review by
globose, their cavities open or closed. Dr Rowan Emberson (Lincoln University) and by Mr
André Larochelle (associate, Landcare Research, Auck-
SYNOPSIS. The carabids, one of the largest beetle fam- land), among others.
ilies in New Zealand, consist of 6 subfamilies (Lawrence Suggested colloquial name: ground beetles (Cicindelinae
& Newton 1995) – CARABINAE, CICINDELINAE, MIGADOPINAE, – tiger beetles).
SCARITINAE, TRECHINAE, and HARPALINAE - with some 445
species in approximately 75 genera and 16 tribes. KEY TO SUBFAMILIES OF CARABIDAE
Many carabids are unique to New Zealand, e.g., the Modified from Lawrence & Britton (1994), following the
endemic genus Mecodema with some 60 species, mostly arrangement of Lawrence & Newton (1995). For identi-
large and robust in form, which apparently occupy similar fication of tribes, consult the key provided as Appendix 3.
ecological niches to the northern temperate genus Cara-
bus. (Several specimens of Carabus were found in Auck- 1 Antennae inserted dorsally on frons; clypeus broader
land in the 1940s, but none is established.) Many carabids than distance between antennal sockets; eyes ex-
are apterous, and often occupy restricted habitats or tremely large and protruding; legs very long and
geographical areas. There are interesting large species of slender; protibia with 2 terminal spurs; lateral pronotal
Megadromus, e.g., M. antarcticus, which is common on carina absent or incomplete posteriorly
the Canterbury Plains in the South Island. Small species of ... (Fig. 102) .. CICINDELINAE
the large genus Bembidion may commonly be found on —Antennae inserted on side of head between eye and
banks of streams, rivers, and lakes. mandibular scrobe; clypeus narrower than distance
A dominant group of terrestrial predators, with some between antennal sockets; eyes moderately large, not
known to be arboreal, mainly nocturnal but some diurnal. or slightly protruding (exception: Scopodes species,
Encountered in a variety of habitats ranging from forest which have large protruding eyes); without other com-
litter, old tree trunks, under stones, and tussock grass to bination of characters ... 2
some wet habitats, including riparian and coastal areas.
Collecting: baited or unbaited pitfall traps, on the 2(1) Metepimeron not visible between posterior edge of
ground and around the base of tree trunks, sifting organic metepisternum and anterior edge of 1st ventrite;
debris, and hand-picking from logs or under stones or mesocoxal cavities open laterally, partly closed by
bark, at night with a head-light. mesepimeron ... 3
—Metepimeron visible as a lobe between metepisternum
RANGE. North ®, South
®, Stewart ® and 1st ventrite; mesocoxal cavities usually closed
Kermadecs ®, ®, Three Kings
Snares
®, Chathams ®, laterally by meeting of sterna ... 4

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3(2) Procoxal cavities open behind; apical segment of All our dytiscids are aquatic and very good swimmers.
maxillary palp more or less expanded and truncate They often occur in small bodies of water such as ponds,
apically; body length more than 17 mm but may be found in streams and rivers, and even under-
... (Fig. 103) .. CARABINAE ground in waterlogged coarse alluvial gravels. They norm-
-Procoxal cavities closed behind; apical segment of ally occur in well vegetated waters, where adults and
maxillary palp slender and fusiform; body length larvae prey on small arthropods. Dytiscids breathe air
usually 17 mm or less ... (Fig. 104) .. MIGADOPINAE trapped under the elytra, and periodically come to the
surface to renew their air supply. Most species fly, and may
4(2) Body pedunculate, with a distinct, narrow waist or migrate to different sites. Of special interest are phreatic
peduncle between prothorax and elytra, the scutellum water beetles of the genera Kuschelydrus and Phreato-
entirely contained within peduncle ... 5 dessus, with species living in underground waters (Ordish
- Body not pedunculate, or if slightly so then scutellum 1976a, 1991).
extending behind peduncle and forming a wedge Collecting: using an aquatic net in small ponds and
between elytral bases ... 6 other open waters, or by light trapping near water. Samp-
ling the phreatic (underground) fauna requires special
5(4) Mesocoxal cavities open laterally, partly closed by techniques (see, e.g., Ordish 1976a, 1991).
mesepimeron ... (Fig. 105) .. SCARITINAE
- Mesocoxal cavities closed laterally by meeting of RANGE. North ®,®,South
Stewart
sterna ... (Fig. 106) .. TRECHINAE (in part) Kermadecs ®, Three Kings O, Chathams ®, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
6(4) Mandible with 1 setiferous puncture in a scrobe along
outer edge ... (Fig. 106) .. TRECHINAE (in part) EXAMPLES. HYDROPORINAE - Kuschelydrus phreaticus
- Mandible without a setiferous puncture in a scrobe Ordish, 1976 (Fig. 108), Liodessus plicatus Sharp, 1882
... (Fίg. 107) .. HARPALINAE (Fig. 109).

SELECTED REFERENCES. Bousquet (1991b), Britton REMARKS. Suggested colloquial name: diving beetles.
(1940, 1941, 1949, 1958, 1959, 1960a, b, 1962, 1963,
1964), Brouerius van Nidek (1965), Butcher (1984), SELECTED REFERENCES. Ordish (1966, 1974, 1976a,
Gourlay (1950), Johns (1974), Lindroth (1976, 1980), b, 1989, 1991), Spangler (1991a).
Moore (1980), Pilgrim (1963), Rivalier (1964), Townsend
(1965), Watt (1961), Wise (1988, 1990).
[4] Family GYRINIDAE
Fig. 110

[3] Family DYTISCIDAE Length 4.3-4.5 mm Tarsal formula 5-5-5
Fig. 108, 109 DIAGNOSIS. Body glossy, boat-shaped, moderately
Length 2.5-27 mm Tarsal formula 5-5-5 convex to slightly flattened. Eyes completely divided into
upper and lower portions by a strip of cuticle. Antennae
DIAGNOSIS. Body elongate-oval, smooth, convex, short, compact, usually clubbed. Middle and hind legs
with dorsal and ventral surfaces of similar convexity, short, with fringes of swimming hairs; forelegs pro-
dark brown, dark olive, or nearly black, often with nounced and raptorial, with procoxae transverse, and
indistinct yellowish spots or margins. Antennae slender their cavities open.
and threadlike. Mandibles strong. Scutellum visible or
not. Procoxae globose, their cavities open or closed; SYNOPSIS. Represented by one species of GYRININAE,
protibiae lacking spines. Hind legs enlarged, modified Gyrinus convexiusculus, self-introduced to New Zealand
for swimming (moving synchronously), with paddle- from Australia. First recorded from New Zealand as
like tarsi bearing a dense fringe of hairs; metacoxae Gyrinus huttoni (1878), the original specimen was col-
large, lacking plates; metatarsi with either 1 or 2 claws. lected by Capt. F.W. Hutton in the Waikato basin in 1873.
Sometimes males with first 3 basal segments of protarsi No other specimens were seen until the late 1970s, when
dilated to form adhesive pads consisting of suction discs, the species was rediscovered in the Waikato basin. Mr
enabling male to hold female during copulation. Ventral Keith Wise (formerly of Auckland Museum) recently
body without a flattened keel. found it in the Ahipara area of Northland. Immigrant
populations appear to establish in New Zealand
SYNOPSIS. Dytiscids are represented in New Zealand sporadically but not widely.
by some 16 species in 11 genera of subfamilies COPE- The species occurs in small peaty lakes and man-made
LATINAE, HYDROPORINAE, COLYMBETINAE, and DYTISCINAE. dams. Collecting: using aquatic nets.

-22-
RANGE. North ®, South ®?, Stewart O cent club; segment preceding club (`cupule') glabrous
Kermadecs O, Three Kings O, Chathams O, Snares O and often embracing the following segment. Maxillary
Aucklands O, CampbelI 0, Antipodes O, Bounties O palps elongate, often longer than antennae. Legs often
dentate or spinose; procoxae large, transverse and pro-
EXAMPLE. GYRININAE - Gyrinus convexiusculus Mac- jecting, their cavities open; metatarsi flattened and
leay, 1871 (Fig. 110). fringed with hairs. Metasternum often extended at rear
into 4 short spines.
REMARKS. Suggested colloquial name: whirligig beetles.
SYNOPSIS. Approximately 75 species of hydrophilid
SELECTED REFERENCES. Ochs (1949), Spangler are recorded in New Zealand, in some 25 genera and 3
(1991b), Wise (1989). subfamilies - HORELOPHINAE, HYDROPHILINAE, and SPHAER
IDIINAE. Horelophinae are endemic to New Zealand; the
type genus Horelophus is a very atypical hydrophilid.
Suborder POLYPHAGA There are two main groups of species, aquatic and
Comprises the vast majority of Coleoptera-95% ac- terrestrial; many New Zealand hydrophilids are terrestrial,
cording to Lawrence & Britton (1991, 1994), whose and are found in leaf litter or carrion. Larvae tend to be
definition we follow here: prothoracic notopleural suture aquatic or semiaquatic, but many species are known from
absent (Fig. 3); prothoracic pleuron fused with trochantin moist, decaying organic matter. A few species are reported
and entirely concealed, forming a cryptopleuron; hind from thermal waters. Adults are usually herbivorous or
wing without an oblongum cell, and with transverse fold scavengers, whereas all known larvae are carnivorous and
never crossing MP (Fig. 51); 1st abdominal sternite not cannibalistic.
divided by metacoxae (Fig. 3), which are usually movable Collecting: netting in aquatic habitats, sifting decaying
and not fused to metasternum; aedeagus usually with a vegetable matter, or light-trapping.
basal piece.
RANGE. North ®,® South . Stewart ®
Kermadecs . Three
® Kings ®, Chathams ®, Snares ®
Series STAPHYLINIFORMIA Aucklands ®, Campbell O, Antipodes ®, Bounties ®
Adults distinguished by reduced wing venation (except in
Hydrophilidae) and a sophisticated type of wing-folding EXAMPLES. HORELOPHINAE - Horelophus walkeri Orch-
mechanism; larvae usually with articulated urogomphi ymont, 1913 (Fig. 111); ΗYDROΡΗILΙΝΑΕ- Enochrus tritus
(Lawrence & Britton 1991, 1994). (Broun, 1880) (Fig. 112); SΡΗΑΕRIDΙΙΝΑE - Rygmodus
tibialis Broun, 1886 (Fig. 113).

Superfamily HYDROPHILOIDEA REMARKS, New Zealand's diverse fauna of Hydro-

Antennae short, with a long scape and a densely pubescent philidae is much in need of revision at all levels. The
3-segmented club (Fig. 26, 27); segment preceding club Sphaeridiinae are under review by Dr M. Hansen (Zoo-
(`cupule') transverse, concave, and in aquatic species used logical Museum, Copenhagen). The terrestrial forms are
for respiration. Hind wings with a radio-median (R-M) probably important indicators of environmental quality.
loop, without a spring mechanism. Procoxae large; tibiae Suggested colloquial name: water scavenger beetles.
dentate or spinose; tarsi 5-5-5, 5-5-4, or 5-4-4, sometimes
appearing 4-4-4 because of vestigial basal segment. Aede- SELECTED REFERENCES. Hansen (1991b, 1995),
agus trilobed (with fused parameres in most Histeridae). Newton (1989), Orchymont (1913), Ordish (1974,
Larva distinct (for details see Lawrence & Britton 1991, 1976b), Spangler (1991d), Todd (1961), Winterbourn
1994). A phylogenetic analysis of Hydrophiloidea based (1968, 1970), Wise (1965, 1973).
on characters of the adult head and the larvae is provided
by Beutel (1994) and Hansen (1991a, b, 1995).
[6] Family HISTERIDAE
Fig. 114, 115
[5] Family HYDROPHILIDAE
Length 1.5-10 mm Tarsal formula 5-5-5 or 5-5-4
Fig. 111-113
Length 2-9 mm

Tarsal formula 4-4-4 or 5-5-5 DIAGNOSIS. Body robust, disc-shaped, oval or nar-
rowly oval to nearly rectangular with rounded angles,
DIAGNOSIS. Body usually oval, moderately to glabrous, moderately convex to flattened, black and
strongly convex, smooth and usually glossy. Antennae strongly glossy. Head deeply inserted. Antennae unique
short, 7-9-segmented, often concealed from above, in form: short and elbowed, with an abrupt, 3-
with a long scape and a 3-segmented, densely pubes- segmented club, often inserted in a cavity or hidden on

-23-
underside of prothorax. Elytra shortened and truncate, [7] Family HYDRAENIDAE
exposing tip of abdomen (usually 2 or 3 apical segments), Fig. 116, 117
with 6 or fewer striae. Procoxae large and transverse,
their cavities open; protibiae expanded, often spiny. Length 1.3-2.6 mm Tarsal formula 5-5-5
DIAGNOSIS. Body narrowly oval or elliptical (broad
SYNOPSIS. There are 29 nominal New Zealand species in species of Meropathus), flattened to slightly convex,
of histerid, of which 6 are adventive. They occupy some 13 small, with scutellum small; generally similar to Hydro-
genera, including one undescribed, in 5 subfamilies - philidae but with 6 or 7 abdominal segments as opposed
ABRAEINAE, SAPRININAÉ, DENDROPHILINAE, TRIBALINAE, and to 5. Maxillary palps elongate, usually longer than
HISTERINAE. Parepierus is the largest genus, containing antennae. Antennae 9-11-segmented (appearing 8-
about one-third of the species, and Reichardtia is an segmented in Meropathus) and with a 2-, 3-, or usually 5-
endemic New Zealand genus. Most species are widely segmented pubescent club. Legs with last tarsomere
distributed, although histerids are infrequently encountered elongate, often longer than all preceding segments com-
in New Zealand. bined, with no distinct bisetose empodium between claws;
Adults and larvae are almost exclusively carnivorous, procoxae transverse, their cavities open or (Hydraena)
and prey on other insects, usually larvae. They may be closed.
commonly found on carrion, decomposing plant materials,
tree wounds, seashore debris, rotting mushrooms, and SYNOPSIS. Thirty-two species of hydraenid are recog-
under bark in galleries of other insects. Kuschel (1990) nised in New Zealand, in 5 genera and 2 subfamilies -
recorded some species in a suburban environment from HYDRAENINAE and OCHTHEBIINAE. Twenty-two species are
compost, lawn clippings, poultry straw, carcasses, fowl recorded from the South Island and subantarctic islands, 4
manure, bush floor litter, and decayed wood. The smaller from the North Island, and 6 occur on both main islands
species are sometimes common around seabird nests. (Ordish 1984). This relatively rich hydraenid fauna offers
Collecting: hand-picking, or sifting organic material an interesting field for ecological studies.
and processing it in Berlese funnels. All the genera except Meropathus live as adults and
larvae in fast-flowing streams. Meropathus species are
RANGE. North ®, South ®, Stewart terrestrial in the coastal or supralittoral zone, occurring in
Kermadecs ®, Three Kings ®, Chathams ®, Snares O porous rocks, littoral moss and grasses, and seabird nests.
Aucklands O, Campbell O, Antipodes O, Bounties O Adults of the aquatic genera all have one or more pairs of
legs modified (Ordish 1984, fig. 12-16) and feed on dead
EXAMPLES. SAPRININAE - Reichardtia pedatrix (Sharp, leaves.
1876) (Fig. 114), Tomogenius latipes (Broun, 1881) (Fig.
115). RANGE. North ®, South ®, Stewart
Kermadecs O, Three Kings O, Chathams . Snares ®
REMARKS. This family is under revision by Dr Masa- Aucklands ®, Campbell ®. Antipodes®®, Bounties O
hiro Ohara of Hokkaido University, Japan.
Suggested colloquial name: hister beetles, pill beetles. EXAMPLES. HYDRAENINAE - Podaena latipalpis Ordish,
1984 (Fig. 116); OCHTHEBIINAE - Meropathus zelandicus
SELECTED REFERENCES. Dahlgren (1968, 1971, Ordish, 1984 (Fig. 117).
1976), Mazur (1984), Newton (1991 c), Wenzel (1944).
REMARKS. Suggested colloquial names: minute moss
beetles, cascade beetles.
Superfamily STAPHYLINOIDEA
Body shape variable, antlike in Scydmaenidae (Fig. 122), SELECTED REFERENCES. Gressitt & Samuelson
short, stout, and wedge-shaped in Pselaphinae (Fig. 126), (1964a), Hansen (1991a), Ordish (1984), Spangler
ovoid in Hydraenidae (Fig. 116, 117) and Agyrtidae (Fig. (1991c).
119) ovoid to nearly spherical in Ptiliidae and Leiodidae
;

(Fig. 118, 120, 121), and elongate, slender, with exposed

flexible abdomen in most Staphylinidae (e.g., Fig. 132- [8] Family PTILIIDAE
138). Adults distinctive in having usually strongly Fig. 118
projecting coxae, metasternum almost always lacking a
Body length 0.6-1 mm Tarsal formula 2-2-2 or 3-3-3
median suture, legs often spinose, elytra usually truncate,
exposing at least one tergite, hind wing with reduced DIAGNOSIS. Body minute, brown to black, broadly
venation and without a radio-medial (R—M) loop, and elongate-oval. Antennae filamentous, bearing long
aedeagus with a reduced phallobase. Larvae characteristic hairs and with a 2- or 3-segmented club. Elytra often
— see Lawrence & Britton (1991, 1994). short, exposing 1 or 2 abdominal segments. Hind wings

—24—
feather-like, fringed with long hairs. Scutellum large. (1982), and are believed to be closely related to Leiodidae
Procoxae transverse or globose, their cavities open; (Newton 1985).
metacoxae variable in shape, often with large coxal plates. Species of Agyrtidae are known from forested areas,
Tarsi usually 2- or 3-segmented but appearing 1-segmen- and are common on carrion and faeces. Adults of `N'.
ted in some species. prolongatus occur in forested areas on carrion, e.g., dead
birds especially at a slightly mummified stage, or may be
SYNOPSIS. In New Zealand there are some 56 species of found in soil crevices on stream banks. Some specimens
ptiliid, including 8 or 9 adventive, in perhaps 13 genera were collected from an abandoned kiwi egg in a burrow.
and 3 subfamilies — PTILIINAE, NANOSELLINAE, and ACRO- An effective method of collecting adults is carrion-baited
TRICHINAE. pitfall traps. Larvae may be seen in association with adults
Common in decaying organic matter, where apparently on carrion.
they feed on fungal spores and hyphae (Lawrence &
Britton 1991, 1994). Collected from garden compost, leaf RANGE. North ®, South ®, Stewart O
litter, moss mat, forest litter, decayed wood, mouldy dung, Kermadecs O, Three Kings O, Chathams O, Snares O
under bark in native forests, amongst litter and tussocks, Aucklands O, Campbell O, Antipodes O, Bounties O
seashore debris including decaying seaweed, and humi-
fied soil (Johnson 1982, Kuschel 1990). Adults are often EXAMPLE. `Necrophilus' prolongatus (Fig. 119).
caught in flight interception traps.
REMARKS. Suggested colloquial name: carrion beetles.
RANGE. North . South ®, Stewart ®
Kermadecs . Three Kings ®,® Chathams ®. Snares ® SELECTED REFERENCES. Newton (1991a).
Aucklands ®, Campbell ®, Antipodes ®. Bounties O

EXAMPLE. PTILIINAE - Notoptenidium lawsoni (Mat- [10] Family LEIODIDAE

thews, 1873) (Fig. 118). [assisted by S.B. Peck and A.F. Newton, Jr]
Fig. 120, 121
REMARKS. Suggested colloquial name: feather-winged
beetles. Body length 0.9-4.3 mm
Tarsal formula 5-5-5, 5-5-4, or 5-4-4
SELECTED REFERENCES. Dybas (1991), Gressitt & DIAGNOSIS. Body broadly oval to nearly spherical,
Samuelson (1964a), Johnson (l975a—c, 1982). slightly to strongly convex, or scydmaenid-like; surface
glabrous and strongly glossy or pubescent, brown to
black. Antennae with a 3-5-segmented club; when club
[9] Family AGYRTIDAE 5-segmented then segment 8 (2nd segment of club)
Fig. 119 usually smaller than adjoining segments (Fig. 24);
Length 9-10 mm

Tarsal formula 5-5-5 distal surface of antennal segments 7, 9, and 10 with
opening of a unique type of sensory vesicle. Elytra often
DIAGNOSIS. Body broadly oval, slightly depressed, striate, and often with a longitudinally impressed line
with sides of pronotum and elytra explanate, brown, along suture. Procoxae globose and projecting; meta-
glabrous and glossy. Antennae 11-segmented, with a 5- coxae close together. Tibiae often expanded and spiny.
segmented pubescent club. Elytra covering entire
abdomen, with 9 or 10 striae. Hind wings well dev- SYNOPSIS. According to unpublished data of G.
eloped. Procoxae conical, slightly transverse basally Kuschel, A. Newton, and S. Peck (pers. comm.) there are
and projecting, their cavities open. Mesotibiae and some 112 species in approximately 25 currently recog-
metatibiae each with a large spur. Tarsomeres and claws nised genera, including 6 or 7 new genera and nearly 50
large. new species. Four subfamilies are represented — CAM'-
ARINAE, with some 69 species, is the largest, followed by
SYNOPSIS. A small, archaic family with fossil records CHOLEVINAE (25), LEIODINAE (17), and COLONINAE (1).
from the mid Jurassic (Newton 1991a), comprising Kuschel (1990) reported collecting species of Leio-
approximately 8 genera in the world fauna. Most agyrtids dinae in the Auckland area from bush-floor litter, amongst
are Holarctic in distribution, with the exception of two ground plants, from grass mats of Oplismenus and
New Zealand species: ` Necrophilus' prolongatus Sharp, Microlaena, and from an agaric in the bush. He recorded
and an undescribed species (Newton 1985). According to Camiarinae collected in the same area from fungi (Auri-
Newton these belong in an undescribed genus, and are cularia polytricha, Ganoderma applanatum), coarse floor
erroneously placed in Necrophilus. Agyrtidae were re- litter, amongst Gahnia plants, under logs and in cracks of
moved from family Silphidae by Lawrence & Newton stream banks, decayed wood, in Ptychomnion moss mats,

— 25 —
and rotten Pinus radiata and Knightia logs. He collected SELECTED REFERENCES. Daffner (1985), Newton
Coloninae in bush litter, cracked stream banks, and (1991b), Szymczakowski (1964, 1966, 1973).
Malaise traps, and collected Cholevinae from carrion.
Newton (1984) reported adults and larvae of Zearagytodes
(Camiarinae) collected in numbers on bracket fungi, with [11] Family SCYDMAENIDAE
the following new host records: Z. maculifer and several [assisted by G. Kuschel]
allied undescribed species on Ganoderma mastoporum,
G. applanatum, and unspecified bracket fungi, all from the
Fig. 122

North Island. Adults and larvae were browsing on the Length 0.5-3.5 mm Tarsal formula 5-5-5
hymenial surface of Ganoderma. Gut contents of both
DIAGNOSIS. Body reddish brown to dark brown, usu-
stages included spores and hyphae. Newton (1984)
ally ant-like, with a distinct `waist' constriction be-
provided some data for the worldwide genus Colon as
tween prothorax and elytra. Head with a distinct neck.
collected in flight traps and by sweeping vegetation, Antennae long, pubescent, with a more or less distinct
especially grasses in wooded areas, and suspects that the 3- or 4-segmented club. Pronotum ovoid or spherical,
genus is associated with hypogeal fungi or moulds.
often with basal foveae, and without carinae. Elytra
Species of Cholevinae, both adults and larvae, may be oval, covering entire abdomen, with rounded humeri;
found on decaying Organic matter, on which they feed: entire dorsal surface usually glossy and with long hairs.
dung, carrion, well decayed soft fungi, forest litter, nests of
Maxillary palps usually enlarged, with apical segment
vertebrates, etc. (Newton 1984). reduced. Legs with enlarged femora. Procoxae trans-
verse and projecting, their cavities open.
RANGE. North ®, South ®, Stewart
Kermadecs O, Three Kings ®, Chathams ®, Snares SYNOPSIS. Watt (1982a) recorded 165 nominal native
Aucklands ®, Campbell ®, Antipodes O, Bounties O species of scydmaenid; 201 native and 1 adventive species
are now represented in NZAC, many of them undescribed.
EXAMPLES. CAMIARINAE - Inocatops elongellus Broun,
They belong to some 11 currently recognised genera of
1917 (Fig. 120); COLONINAE- Colon hirtale (Broun, 1880) SCYDMAENINAE, of which 6 are endemic to New Zealand.
(Fig. 121).
For classification see Franz (1975, 1977, 1980, 1985).
Kuschel (1990) recorded several species from the
REMARKS. Suggested colloquial name: small carrion Auckland area collected in leaf litter, from decayed wood
beetles. of various trees and hollow logs, moss mat, leaf mould in
tree hollows, and Malaise traps. Scydmaenids are noc-
KEY TO SUBFAMILIES OF LEIODIDAE turnal, living in concealed habitats, with some species
IN NEW ZEALAND [by A.F. Newton, Jr] flying at dusk. Good collections might be obtained by
1 Head with an elevated crest (occipital carina) along processing sifting organic litter through Berlese funnels.
hind margin, resting against front of pronotum when
head in repose ... CHOLEVINAE RANGE. North ®, South ®, Stewart ®
—Head without an elevated crest along hind margin .. 2 Kermadecs O, Three Kings ®, Chathams ®, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
2(1) Antennae 11-segmented, with segment 8 as large as
segments 9 and 10, and club of 4 segments (Fig. 25); EXAMPLE. SCYDMAENINAE - Adrastia clarkei (Franz,
dorsum setose ... (Fig. 121) .. COLONINAE 1985) (Fig. 122).
—Antennae usually 11-segmented, with segment 8
smaller than segments 7 and 9, and club usually of 5 REMARKS. Suggested colloquial name: stone beetles.
segments (Fig. 24); if club 4-segmented then antennae
10-segmented and dorsum glabrous ... 3 SELECTED REFERENCES. Franz (1975, 1977, 1980,
1985).
3(2) Antennal insertions concealed in dorsal view; head
relatively flattened and broad, usually half or more as
wide as pronotum; dorsum apparently glabrous
(sometimes with many short hairs visible only at [12] Family STAPHYLINIDAE
higher magnification) ... LEIODINAE (incl. PSELAPHINAE, SCAPHIDIINAE)
—Antennal insertions exposed in dorsal view; head [assisted by A.F. Newton and M.K. Thayer]
relatively convex and narrow, usually less than half as Fig. 123-138
wide as pronotum; dorsum usually with evident long
setae (exceptions: Asphaerites, Catopsolius, Chela-

Length 0.6-25 mm Tarsal formula 5-5-5
gyrtodes) ... (Fig. 120) .. CAMIARINAE or heteromerous

-26-
DIAGNOSIS. Body brown to black, less frequently vegetation. The majority of known species are predators,
brightly coloured or metallic. Form very diverse, usually feeding on larvae and adults of other insects including
narrowly elongate, approximately subparallel, rarely Collembola, and on mites and nematodes.
short, oblong, or wedge-shaped, or short, stout, and oval Species of Osoriinae and Oxytelinae feed on decom-
with abdomen non-flexible (Pselaphinae, Fig. 126). Head posing organic matter (McColl 1982, Newton 1990).
often with a posterior constriction, sometimes with ocelli Some Tachyporinae and Aleocharinae (e.g., Gyrophaena)
(Omaliinae, Microsilphinae). Antennae with 10 or 11 feed on moulds or other fungi. Species of the aleocharine
segments, filiform, beadlike; last 1-4 segments enlarged genus Myllaena were recorded feeding on algae (Klima-
(Pselaphinae), incrassate, or clubbed. Elytra nearly szewski 1992). Larvae of the ectoparasitic genus Aleo-
always short and truncate, exposing a variable number chara (Aleocharinae) are exclusive parasitoids of puparia
of abdominal segments, usually 6 or 7, exceptionally of cyclorrhaphous Diptera. The first-instar larva searches
concealing alI or most of abdomen (Microsilphinae and for a host pupa, gnaws a hole in the puparium, crawls
some Omaliinae, Proteininae, Scaphiidinae) or leaving at inside, seals the entrance behind, and feeds on the pupa
least 2 (usually 3 or 4) tergites exposed (Pselaphinae). (Klimaszewski & Crosby 1997; Klimaszewski & Jansen
Procoxae usually conical and strongly projecting, their 1993).
cavities usually closed. Abdomen flexible dorsoventrally The majority of adult pselaphines occur in leaf and
(exception: Pselaphinae), well sclerotised except basal wood litter on the forest floor. Newton & Chandler (1989)
tergite or two. reported that in general pselaphine species can also be
discovered in wetland, grassland, desert, beach, cave, and
SYNOPSIS. Staphylinids are one of the largest and most even arboreal habitats, wherever debris, moss, or root mats
diverse families of beetles, as of 1987 comprising world- maintain microhabitats of high humidity. Kuschel (1990)
wide about 42 000 nominal species (8400 species of collected pselaphines in Auckland from decayed wood on
Pselaphinae) in several thousand genera (1091 genera in the beach and amongst low coastal vegetation, coastal
Pselaphinae) and 100 tribes (Newton 1985, 1990, Newton litter, hollow trees, wood mould, stream litter and stream
& Chandler 1989, Newton & Thayer 1992). Hundreds of banks, moss, sedges, grasses, soil around tree stumps, etc.
new species are described every year. We follow Newton Adults and larvae of pselaphines are predators, feeding on
& Thayer (1995) and Lawrence & Newton (1995) in mites, springtails, and other small invertebrates.
treating the former family Pselaphidae as a subfamily of More details on the natural history of New Zealand
Staphylinidae. A revised list of the subfamilies, tribes, and staphylinids are provided by Klimaszewski et al. (1996).
genera with synonyms is provided by Klimaszewski et al. Collecting: hand-collecting with the aid of an aspirator;
(1996). pitfall traps, unbaited or baited with carrion, faeces, or
In New Zealand there are approximately 766 nominal fermented fruits; processing sifted organic litter through
native species and 85 adventive, plus 170 or more recog- Berlese funnels or Winkler/Moczarski extractors; Malaise
nised but not yet named. The number of New Zealand traps, window traps, and other flight interception traps.
native staphylinids may exceed 1000 (G. Kuschel, pers.
comm.), making it after Curculionidae our second largest RANGE. North ® , South ®, Stewart
group of beetles. Our species are grouped in some 190 Kermadecs ®, Three Kings ®, Chathams ®, Snares ®
genera and 16 subfamilies, as follows: ALEOCHARINAE 61 Aucklands ®. Campbell ®, Antipodes ®, Bounties Ο
genera (probably an underestimate); EUAESTHETINAE 4;
HABROCERINAE 1; MICROSILPHINAE 1; OMALIINAE 16, and 4 EXAMPLES. ΜICROSILPHINAE — Microsilpha litorea
undescribed; OSORIINAE 3 (including one of former Broun, 1886 (Fig. 123); OMALIINAE — Omaliomimus
Eleusiinae); OXYTELINAE 7; PAEDERINAE 10; PHLOEOCHARINAE albipennis (Kiesenwetter, 1877) (Fig. 124); ΡRΟΤΕΙΝΙΝΑE
2; PIESTINAE 1; PROTEININAE 4; PSELAPHINAE 41, and 20 nitidus Broun, 1895 (Fig. 125); PSELAPHINAE —-Silphoteus
undescribed; PSEUDOPSINAE 1; SCAPHIDIINAE 5; STAPHYLININAE Sagola laminata Broun, 1893 (Fig. 126); ΡHLOEOCHARINAE
26; and TACHYPORINAE 3 (Klimaszewski et al. 1996; Watt — Pseudophloeocharis australis (Fauvel, 1900) (Fig.
& McColl in McColl 1982, updated). 127); ΤACHYPORINAE — Sepedophilus sp. (Fig. 128);
Staphylinid beetles are very successful and broadly HABROCERINAE Habrocerus capillaricornis (Gravenhorst,
—

distributed, occurring in almost all terrestrial habitats. 1806) (Fig. 129); ΑLEOCHARINΑE —hAalemocnrdi
Staphylinids may be found in decomposing organic matter (Klimaszewski & Crosby, 1997) (Fig. 130); SCAPHIDIINAE
such as garden and forest litter, mushrooms, bird and — `Baeocera' scutellaris (Redtenbacher, 1867) (Fig. 131);
mammal nests, decaying seaweeds and other algae of tidal ΡIΕSTIΝΑΕ—P(1aCrFs9imgone4uhd,.)
zones, uncompacted soil, organic matter in tree hollows 132); OSORIINAE — Nototorchus ferrugineus (Broun, 1893)
and burrows, under bark, in galleries of other insects, wet (Fig. 133); OXYTELINAE — Carpelimus sp. (Fig. 134);
moss, cracks on edges of streams, shores of lakes, etc. ΕUΑΕSΤΗΕΤΙΝΑΕ — Agnosthaetus vicinus (Broun, 1921)
Some species may be found on carrion and in animal (Fig. 135); ΡSEUDOPSINAE — Pseudopsis arrowi Bernhauer,
droppings, and some Omaliinae have been beaten from 1939 (Fig. 136); PAEDERINAE — Medon zeelandicus

-27-
(Redtenbacher, 1867) (Fig. 137); STAPHYLININAE — Cafius elytra, with head and pronotum narrower; setose
litoreus Broun, 1880 (Fig. 138). foveae present on head, prothorax, and other parts of
body; elytra short, closely applied to body, leaving
REMARKS. Staphylinids are an ancient group with fossil much of abdomen exposed; abdomen broad, not
records from the early Jurassic period (Newton 1990). The flexible; maxillary palps often modified; antennae
New Zealand fauna of staphylinids may be characterised often beadlike; tarsi 3-segmented
by (1) the absence of several subfamilies from the native ... (Fig. 126) .. PSELAPHINAE
fauna, a few of which are, however, represented by ad- —Body not so; setose foveae absent; elytra usually
ventive species (e.g., Habrocerinae); (2) large species loosely applied to body; abdomen flexible; maxillary
radiations in Eupines, Euplectopsis, Hyperomma, Micro- palps usually simple; antennae not beadlike; tarsi
silpha, Ocalea complex, Oligota, Othius, Paratorchus, usually of more than 3 segments ... 5
`Quedius', Sagola, and Sepedophilus; and (3) high
endemism — approx. 43% at the generic level, even higher 5(4) Abdomen without paratergites; tergites and sternites
at the species level (Klimaszewski et al. 1996); (4) strong usually fused together; body either cylindrical or
southern connections — over 20 groups or genera are strongly flattened ... (Fig. 133) .. OSORIINAE
shared only with other southern temperate regions (New- —Abdomen with paratergites; tergites and sternites of
ton 1985). In general our staphylinids are poorly known; abdomen not fused together; body not cylindrical .. 6
the Aleocharinae, as everywhere, are the least known
amongst staphylinids, and consequently are in urgent need 6(5) Head with paired ocelli on vertex between posterior
of revision. The classification of Staphylinidae at all margins of eyes; paratergites single
levels, including family limits, is unsatisfactory and un- ... (Fig. 124) .. OMALIINAE
stable (Newton 1990). —Head without ocelli (sometimes with a pair of de-
Suggested colloquial name: rove beetles. pressions on vertex); paratergites usually double ... 7

7(6) Antennal insertions located between eyes and

KEY TO SUBFAMILIES OF STAPHYLINIDAE posterior to a line drawn between anterior edges of
OCCURRING IN NEW ZEALAND eyes (except in forms with reduced eyes); terminal
[Modified from Watt & McColl in McColl (1982) to segment of maxillary palp usually very small and
include Microsilphinae, Pselaphinae, and Scaphidiinae.] sharp-pointed ... (Fig. 130) .. ΑLΕΟCΗARΙΝE
—Antennal insertions located anterior to a line drawn
1 Antennae distinctly clubbed (e.g., Fig. 123, 135); body between anterior edges of eyes, or inserted under
without setose foveae ... 2 shelf-like dorsolateral margins of frons; terminal
—Antennae usually not distinctly clubbed; if so, body segment of maxillary palp usually not as above ... 8
compact and with setose foveae, and tarsi 3-segmented
(e.g., Fig. 124-134) ... 3 8(7) Body strongly depressed; antennae inserted under
widely explanate dorsolateral projections of frons,
2(1) Antennal club with more than 2 segments; elytra their sockets facing ventrally; procoxae globular
elongate, covering most of abdomen; anterior margin ... (Fig. 132) .. PIESTINAE
of labrum not as below; body short, oval; head with —Body rarely strongly depressed; antennae usually not
paired ocelli on vertex... (Fig. 123) .. MICROSILPHINAE as above, but if inserted under dorsolateral projections
—Antennae with a 2-segmented club; elytra short, ex- of frons then these are less prominent, and their
posing entire abdomen; anterior margin of labrum sockets face laterally; procoxae elongate, transverse,
minutely crenulate or denticulate; body narrow, elon- or projecting ... 9
gate; head without ocelli.. (Fig. 135) .. EUASTHIN
9(8) Abdomen usually strongly flattened and broad, with 7
3(1) Body wedge-shaped, short, convex and strongly visible sternites, excluding genital segment (exception:
glossy; elytra truncate posteriorly, concealing all but 1 Coprostygnus, with 6 obvious sternites but also with
or 2 abdominal tergites, and exposing pointed abdo- elytral striae, which are rare in taxa keying to the
men; legs and antennae long and slender alternative, below); gular sutures confluent, or at least
... (Fig. 131) .. SCAPHIDIINAE partially so ... (Fig. 134) .. OXYTELINAE
—Body not so; elytra exposing at least 3 abdominal —Abdomen usually less flattened and moderately broad,
tergites (exception: Silphotelus, Proteininae, Fig. with 6 visible sternites (Fig. 124, 127) excluding
125); legs and antennae usually moderately elongate genital segment; gular sutures separate ... 10
and stout ... 4
10(9) Elytra elongate, leaving no more than 5 abdominal
4(2) Body compact, usually widest at abdomen or tip of tergites exposed (in Silphotelus elytra almost com-

— 28 —
 pletely covering abdomen); lateral margin of frons Series SCARABAEIFORMIA
with a small, semicircular emargination on either side, Superfamily SCARABAEOIDEA
just in front of eye; body length usually less than 2 mm
(maximum 4 mm) ... (Fig. 125) .. PROTEININAE Characterised by a highly modified prothorax with large
-Elytra shorter, leaving at least 6 abdominal tergites coxae, usually dentate tibiae with a single apical spur,
exposed; lateral margin of frons not so ... 11 lamellate antennal club (Fig. 30), no metacoxal plates, 2nd
abdominal sternite represented by a lateral portion only,
11(10) Abdominal tergites except the last each with a tergite 8 forming a pygidium, hind wing with reduced
diagonal impressed line on either side; pronotum and venation and with a spring mechanism for folding the
elytra longitudinally costate; tarsi short, about one- wing, and larvae grub-like and usually C-shaped (Fig. 98)
third as long as tibiae; body length 3 mm (Lawrence & Britton 1991).
... (Fig. 136) .. PSEUDOPSINAE
-Abdominal tergites without such lines; pronotum and
elytra not longitudinally costate; tarsi usually more [13] Family LUCANIDAE
than one-third as long as tibiae ... 12 [assisted by B.A. Holloway]
Fig. 139-143
12(11) Head with sides converging evenly posteriorly, Length 6-30 mm Tarsal formula 5-5-5
without a neck constriction clearly visible from above
(Fig. 127-129) ... 13 DIAGNOSIS. Body robust, medium-sized to large,
- Head constricted behind eyes to form a distinct neck, brown to black, often with paler spots forming a pattern,
clearly visible from above (Fig. 137, 138) ... 15 either glabrous and glossy or clothed with scales. Head
prognathous, with well developed mandibles which are
13(12) Elytra with epipleura not delimited, their sides often sexually dimorphic - pronounced in males, and
completely rounded off; body shape distinctive, as bearing teeth or larger processes. Antennae distinct, 10-
figured ... (Fig. 127) .. PHLOEOCHARINAE segmented, usually elbowed, with a comb-like club
- Elytra with epipleura clearly delimited by a carina.. 14 consisting of 3 or 4 thick, lamellate segments that
cannot be held together (Fig. 30). Pronotum usually
14(13) Antennae stouter, expanded towards apex; head short. Abdomen with 5 ventrites.
and pronotum usually densely pubescent; body shape
distinctive, as figured ... (Fig. 128) .. TACHYPORINAE SYNOPSIS. Holloway (1961, 1962, 1963b) recognised
-Antennae very slender, filiform (Fig. 18), with long 26 species in 4 genera. There are also 3 adventive species
tactile setae; head and pronotum glabrous except for from Australia, each in a separate genus (Holloway 1997).
some long tactile setae; body shape distinctive, as Four subfamilies are represented - AESALINAE, SYNDESINAE,
figured ... (Fig. 129) .. HABROCERINAE LAMPRIMINAE, and LUCANINAE.
The species are approximately evenly divided between
15(12) Maxillary palp with apical segment always re- the North and South islands, with two endemic species
duced, less than half as long as penultimate segment; restricted to the Chatham Islands and one to the Moko-
antennal insertion obscured by a dorsolateral shelf- hinau Islands (Holloway 1961). Approximately half the
like expansion of margin of frons species are flightless and half fully winged. The flightless
... (Fig. 137) .. PAEDERINAE species are often allopatric, with restricted ranges in
- Maxillary palp with apical segment variable in length, lowland areas, whereas fully winged species tend to have
but at least half as long as penultimate segment; broad and overlapping ranges, though some are restricted
antennal insertions exposed to montane areas (Holloway 1963a).
... (Fig. 138) .. STAPHYLININAE Hudson (1934) reported lucanids as being mainly noc-
turnal, the adults feeding on tree sap, and larvae living on
SELECTED REFERENCES. Bernhauer (1939, 1941, old decaying trees or their roots, and with a life cycle of
1943), Cameron (1944, 1945, 1946, 1947, 1950), Fauvel several years. Holloway (1963a) suspects that all adult
(1900), Frank (1991), Hammond (1975), Herman (1975), lucanids feed on sap, and are not plant-specific, the larvae
Kasule (1966, 1968, 1970), Klimaszewski (1979), Klima- completing their development in rotten wood or soil. It is
szewski & Crosby (1997), Klimaszewski et al. (1996), widely accepted that the larvae feed on rotten wood or
Leschen & Löbl (1995), McColl (1982, 1984a, b), Newton other decomposing organic material. Ceratognathus pas-
(1984, 1985, 1989, 1990), Newton & Chandler (1989), saliformis is almost certainly myrmecophilous, found in
Newton & Thayer (1992, 1995), Park & Pearce (1962), endemic formicine ant nests (Holloway 1962). Hudson
Sharp (1876a, b), Steel (1949, 1950a, b, 1953, 1960, 1964, (1934) reported Ceratognathus gibbosus as being beaten
1966, 1970), Thayer (1985), Williams (1976). from shrubs in hot sunshine, from December until March.

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RANGE. North ®, South ®, Stewart ® [15j
[15] Family SCARABAEIDAE
Kermadecs ®. Three Kings ®, Chathams ®,® Snares O [assisted by R.M. Emberson]
Aucklands O, Campbell O, Antipodes O, Bounties O
Fig. 145-152
EXAMPLES. AESALINAE - Ceratognathus parrianus Length 3-25 mm Tarsal formula 5-5-5
(Westwood, 1863) (Fig. 139); LUCANINAE - Paralissotes
reticulatus (Westwood, 1847), male (Fig. 140), Geodorcus DIAGNOSIS. Stout-bodied beetles, often glossy but
auriculatus (Broun, 1903), male and female (Fig. 141,
some dull (e.g., Saprosites, Aphodiinae), glabrous or
142); LAMPRIMINAE - Dendroblax earlii (White, 1846) moderately to densely pubescent, usually brown or black
(Fig. 143). but occasionally brightly coloured (some Melolonthinae).
Either labrum or mandibles, or both, concealed from
above by clypeus. Antennae 8-10-segmented, not
REMARKS. This family is under revision by Dr B.A. elbowed; club variable, usually 3-, 4-, or 5-segmented
Holloway, of Auckland. but not cupuliform; lamellate segments appearing as
Suggested colloquial name: stag beetles. elongate or oval lobes that can be folded together
tightly to form a compact and asymmetrical club.
SELECTED REFERENCES. Carlson (1991a), Holloway Sculpture of elytra not rough. Legs strongly modified for
(1960, 1961, 1962, 1963a, b, 1996, 1997). digging with teeth, spines, and/or bristles. Abdomen
with 6 ventrites.

[14] Family TROGIDAE SYNOPSIS. Watt (1982a) recorded approximately 132

Fig. 144 native and 12 adventive species of scarabaeid in New
Zealand. This relatively large group of beetles represents 4
Length 6.4-6.5 mm Tarsal formula 5-5-5 subfamilies—MELOLONTHINAE, SCARABAEINAE, DΥΝΑSΤΙΝΑE,
APHODIINAE—and some 25 genera, 13 of them endemic.
DIAGNOSIS. Body robust, oval, slightly subparallel, The subfamily Melolonthinae is by far the largest, with
heavily sclerotised, with dorsal surface rugose, lus- 103 valid native species in 10 genera (Given 1952, 1954,
trous, and often covered with debris. Head strongly 1960 (key to genera), 1966, Watt 1984, Emerson & Barratt
deflexed. Antennae 10-segmented, elbowed, with a 1997). Species of Prodontria are all flightless, have
compact lamellate club (Fig. 31). Procoxae projecting; limited ranges, and are vulnerable to habitat modification.
mesocoxal cavities broadly closed. The Cromwell chafer, P. lewisi, is considered to be an
endangered species, and a nature reserve has been est-
SYNOPSIS. New Zealand has only a single adventive ablished near Cromwell to preserve it (Watt 1984). The
European species of trogid. This was first found at Kumeu, endemic manuka beetle, Pyronota festiva, is common all
north-west of Auckland, in deep litter of poultry houses in over New Zealand, and was first collected during Captain
association with mummified rat carcasses. Other records Cook's voyage in 1769. The 9 greenish-coloured species
include Campbell's Beach near Matakana, Western Hills of Stethaspis are common in forest habitats. This sub-
near Whangarei, the Noises Islands, and Hastings. family contains many species which are recognised pests
Adults have been collected near rat carcasses, under the of agriculture, e.g., the grass grub Costelytra zealandica.
hide of a long-dead cow, in dry animal remains, in wood The Scarabaeinae, with some 14 species, include native
mould in oaks and elms, and in birds' nests (Brown 1967). species in 2 endemic genera and 4 adventive species in 3
Larvae of other species are known to occur in vertical exotic genera (Emberson & Matthews 1973, Paulian 1935,
burrows beneath carcasses (Carlson 1991b, Lawrence & Watt 1984). The Dynastinae include some 5 native species
Britton 1991, 1994). in the endemic genus Pericoptus (Endrödi 1974, Watt
1984) and 3 adventive species in exotic genera (Watt
RANGE. North . South O, Stewart O 1984). The Aphodiinae are represented by 6 genera, with
Kermadecs O, Three Kings O, Chathams O, Snares O some 10 native species and several adventive (Richards
Aucklands O, Campbell O, Antipodes O, Bounties O 1959, Watt 1984).
Scarabaeids include scavenging and/or phytophagous
EXAMPLE. Trox scaber (Linnaeus, 1767) (Fig. 144). species. The melolonthines have species occurring in
many different terrestrial habitats: sand dunes, manuka
(Leptospermum scoparium) scrub, mixed scrub and forest
REMARKS. Suggested colloquial name: carcass beetles. margin, northern and southern rain forest, subalpine scrub
and tussock, river or lakeside sands, and pasturelands
SELECTED REFERENCES. Britton (1956), Brown (Given 1952). Many species are economically important,
(1967), Carlson (1991b), Scholtz (1982, 1986). with larvae feeding on plant roots (Hoy & Given 1952).

-30--
30-
The aphodiines are mainly dung feeders and general Matthews (1973), Emerson & Barratt (1997), Emerson &
® to sandy
scavengers. The endemic dynastines are confined Wallis (1994), Endrödi (1974), Given (1952, 1954, 1960,
seashores and river floodplains (Given 1955). The en- 1964, 1966), Hoy & Given (1952), Lowe (1961), Richards
demic scarabaeines occur predominantly in forest leaf (1959), Scholtz (1990), Stebnicka & Howden (1995),
litter, and probably feed on dung of bats, birds, reptiles, Watt (1971, 1979a, 1984).
and large snails (Emberson & Matthews 1973).

RANGE. North ®, South ®, Stewart ® Series ELATERIFORMIA

Kermadecs . Three Kings ®, Chathams ®, Snares ® For our concept of this series we have followed Lawrence
Aucklands O, Campbell O, Antipodes O, Bounties O & Newton (1995), who define it in Crowson's (1960)
original sense, with Scirtoidea and Dascilloidea included
EXAMPLES. APHODIINAE -Acrossidius tasmaniae as basal lineages. This series in New Zealand includes four
(Hope, 1847) (Fig. 145), Phycochus graniceps Broun, superfamilies: Scirtoidea, Buprestoidea, Byrrhoidea, and
1886 (Fig. 146); SCARABAEINAE Onthophagus granulatus
-
Elateroidea. Phylogenetic relationships among the super-
Boheman, 1858 (Fig. 147); MELOLONTHINAE Odontria-
families are still not well understood (Lawrence & Newton
giveni Watt, 1984 (Fig. 148), Costelytra zealandica 1995). A cladistic analysis of this group was recently
(White, 1846) (Fig. 149), Prodontria lewisi Broun, 1904 conducted by Lawrence et al. (1995).
(Fig. 150); DYNASTINAE Heteronychus arator (Fabricius,
-
Lawrence & Britton (1991, 1994) excluded Scirtidae,
1775) (Fig. 151), Pericoptus truncatus Fabricius, 1775 Eucinetidae, and Clambidae from Elateriformia, and
(Fig. 152). defined the group by the following major characters:
heterogeneous life cycle, with long-lived larvae and short-
REMARKS. Suggested colloquial names: scarab beetles, lived adults usually occupying different habitats; surface-
dung beetles, chafer beetles. active adults often with body streamlined and with a
complex pro-mesothoracic interlocking device or, in its
absence, a chemical defence system; antennae rarely
KEY TO SUBFAMILIES OF SCARABAEIDAE clubbed, often serrate; metacoxae usually excavated to
1 Head with anterior margin more or less semicircular, receive femora.
or emarginate medially; clypeal carina continuous
with canthus, which extends further laterally than eye;
mesocoxae strongly oblique; antennae with 8 or 9 Superfamily SCIRTOIDEA
segments ... 2 Comprises three families with a compacting mechanism in
- Head with anterior margin not as above; clypeal carina which the head is strongly hypognathous and fits against
separated from canthus by a pronounced emargination the procoxae or metasternum in the resting position
on either side; mesocoxae transverse or slightly obli- (Lawrence & Britton 1991, 1994, q.v. for larval char-
que; antennae often 10-segmented ... 3 acteristics).
2(1) Metatibiae each with 2 apical spurs; mesocoxae
oblique and approximated; elytra usually concealing
entire abdomen; scutellum present; body usually sub- [16] Family SCIRTIDAE
parallel ... (Fig. 145, 146) .. APHODIINAE (=CYPHONIDAE =DASCYLLIDAE =DISCYLLIDAE =HELODIDAE)
- Metatibiae each with 1 apical spur; mesocoxae distally Fig. 153, 154
separated; elytra shortened, leaving pygidium ex- Length 1.5-10 mm Tarsal formula 5-5-5
posed; scutellum not apparent; body stout, oval or
rounded ... (Fig. 147) .. SCARABAEINAE DIAGNOSIS. Body brown to black, narrowly oval to
nearly round in outline. Head narrow, with sharp genal
3(1) Head and pronotum in males simple; labrum sclero- ridges which rest against the procoxae when head is
tised, exposed; mandibles concealed from above fully deflexed (Fig. 12); eyes smalI but prominent.
... (Fig. 148-150) .. MELOLONTHINAE Antennae filiform. Pronotum short and transverse,
- Head and pronotum in males usually with horns, often concealing head, covered by fine, recumbent
tubercles, or complex elevations; labrum membranous, pubescence. Elytra often with ridges, and with com-
concealed; mandibles usually partly visible from plete epipleura. Procoxae large and projecting, with
above ... (Fig. 151, 152) .. DYNASTINAE prosternal region reduced; metacoxal plates often
present; 4th tarsal segment lobed beneath.
SELECTED REFERENCES. Bain (1980), Barratt
(1982), Barratt & Campbell (1982), Brown (1967), SYNOPSIS. Scirtids are well represented in the New
Carlson (1991b), Dymock & Forgie (1993), Emberson & Zealand fauna, and in the southern temperate region in

-31-
general. Watt (1982a) recorded 121 nominal native spe- RANGE. North ®, South ®, Stewart O
cies. Some 125 species are housed in NZAC, and are Kermadecs O, Three Kings O, Chathams O, Snares O
placed in 11 genera. Scirtids are poorly known in New Aucklands O, Campbell O, Antipodes O, Bounties O
Zealand; ` Cyphon' particularly is a widespread genus in
need of revision. EXAMPLE. Eucinetus stewarti (Broun, 1881) (Fίg.
Adults may be quite common on vegetation near water, 155).
and on flowers, and larvae of some species are associated
with water in the base of epiphytic plants, e.g., Collo- REMARKS. Suggested colloquial name: plate-thigh
spermum hastatum, particularly in wet forest areas. Larvae beetles.
of Veronatus and its allies live in humus-rich soil. In
general, scirtid larvae are considered to be filter-feeding
detritivores (Beier 1952). SELECTED REFERENCES. Vit (1977a, b), Wheeler &
Collecting: beating shrubs, sweeping vegetation in wet Hoebeke (1984).
places, sifting organic matter near water, intercept traps,
and light traps.
[18] Family CLAMBIDAE
RANGE. North ®, South ®, Stewart ® Fig. 156
Kermadecs O, Three Kings ®, Chathams ®, Snares O Length 0.8-1.4 mm Tarsal formula 4-4-4
Aucklands O, Campbell O, Antipodes O, Bounties O
DIAGNOSIS. Body minute, globular, strongly glossy,
with sparse and indistinct pubescence; many species are
EXAMPLES. Amplectopus pallicornis Broun, 1886 (Fig. capable of partially rolling themselves into a ball. Head
153), Veronatus tricostellus White, 1846 (Fig. 154). strongly deflexed, large, as broad as three-quarters of
maximum pronotal width. Clypeus pronounced. Eye
REMARKS. Sharp (1878) divided the New Zealand simple, or divided by a genal canthus into dorsal and
scirtids into two groups, those with a distinct antennal ventral halves (genus Clambus). Antennae 10-segmented,
fossa and those without a fossa or with a weak fossa. with a 2-segmented club. Scutellum triangular, large.
Suggested colloquial name: marsh beetles. Procoxae projecting, their cavities open; metacoxae
with expanded plates concealing ventrite 1 and folded
SELECTED REFERENCES. Hannappel & Paulus legs. Legs short and slender.
(1992), LeSage (1991a).
SYNOPSIS. Watt (1982a) reported 5 native and 2
adventive nominal species of clambid from New Zealand.
[17] Family EUCINETIDAE Endrödy-Younga (1990) revised the New Zealand clam-
Fig. 155 bids, recording 11 species grouped in 2 genera.
Clambids may be found in leaf litter, lawn clippings,
Length 3.0-3.6 mm Tarsal formula 5-5-5 garden prunings, and compost. Some species are attracted
DIAGNOSIS. Body streamlined, elliptical, uniformly to light, and can be collected using a UV light trap.
brownish to black, with fine decumbent pubescence. Lawrence & Britton (1991, 1994) reported adults feeding
Head deflexed, concealed from above and resting on fungal spores. Kuschel (1990) collected species of
against procoxae. Antennae filiform. Eyes small but Sphaerothorax in the Auckland area on tree stumps and in
prominent. Prothorax reduced. Elytra tapering post- loose moss mats of Polytrichaceae, in organic litter, in a
eriorly, often with fine cross-striations and a longi- heap of Eucalyptus branches and cut bamboo in a sheep
tudinal impressed line on either side of suture. paddock, in a litter sample taken in Pittosporum and
Procoxae projecting, their cavities open; metacoxae Melicytus scrub, and at the foot of a high coastal cliff.
expanded, with oblique metacoxal plates enlarged and
partly concealing 1st ventrite. Legs short; tibiae and tarsi RANGE. North ®, South ®, Stewart ®
of middle and hind legs bearing combs of dark spines. Kermadecs O, Three Kings ®, Chathams O, Snares O
Adults are capable of jumping using their modified hind Aucklands O, Campbell O, Antipodes O, Bounties O
legs.
EXAMPLE. Clambus domesticus Broun, 1886 (Fig. 156).
SYNOPSIS. Eucinetus stewarti, a native species, is the
only New Zealand eucinetid, and little is known concern-
ing its status and habits. Northern temperate species are REMARKS. Suggested colloquial name: clam beetles.
known to feed on fruiting bodies of basidiomycete fungi or
on slime mould spores (Wheeler & Hoebeke 1984). SELECTED REFERENCES. Endrödy-Younga (1990).

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Superfamily BUPRESTOIDEA SELECTED REFERENCES. Bellamy & Williams
The phylogenetic affiliations of Buprestidae are not (1985), Clark (1938), Dumbleton (1932), Milligan (1974,
resolved. The group is currently placed in a monotypic 1985), Morgan (1966), Stuart (1955).
superfamily (Lawrence & Britton 1991, 1994). Lawrence
(1988) suggested that Buprestidae could be the sister-
group of the Byrrhoidea, or might be derived from a Superfamily BYRRHOIDEA
dascilloid ancestor. The superfamily concept used here is after Lawrence &
Britton (1991). Almost all species of Byrrhoidea are
aquatic, semiaquatic, or associated with foliage beside
[19] Family BUPRESTIDAE streams. They have 5-5-5-segmented tarsi with the en-
larged last segment bearing pronounced claws (except for
Fig. 157 Heteroceridae and some Byrrhidae, which have 4-4-4
Length 1.7-10 mm Tarsal formula 5-5-5 segmented tarsi). Adults have a sophisticated type of
DIAGNOSIS. Body bullet-shaped, subparallel, pointed wing-folding mechanism, and larvae usually have a single
at elytral tip, metallic green or black, often with bright pretarsal seta.
yellow spots, more or less glossy, heavily sclerotised, Adults of Byrrhidae, Dryopidae, Elmidae, Limnichidae,
appearing glabrous or with scattered hairs. Head strongly and Ptilodactylidae have the following characteristics:
deflexed, deeply inserted into prothorax. Prothorax head more or less concealed by pronotum from above; tarsi
closely applied to elytra. Antennae moderately long, 5-5-5-segmented, with last segment enlarged; legs re-
threadlike to slightly serrate. Mesosternum with a large tractable into body cavities; prosternum with a process
cavity for reception of prosternal process; metasternum fitting into a cavity of mesosternum (Fig. 52, 53); body
with a curved transverse suture. First 2 abdominal convex dorsally; antennae slender, sometimes weakly or
sternites strongly connate, the suture between them distinctly clubbed. The family Heteroceridae is the most
vaguely defined. Procoxae globose, their cavities open. divergent in this group, and distinguished by prognathous
Tarsi with segments 1-4 lobed below. head, short, thick, sawtoothed antennae, flat, spinose
tibiae, and 4-4-4-segmented tarsi, with the last segment
SYNOPSIS. Watt (1982a) recorded a total of 4 nominal not pronounced. The family Chelonariidae is also quite
species of buprestid, whereas Dumbleton (1932) and distinct, but probably close to the main byrrhid lineage.
Clark (1938) had indicated 2 native species and 1 adven-
tive. The native species belong to 2 genera of BUPRESTINAE.
Adults and larvae of Nascioides enysi are found in [20] Family BYRRHIDAE
Nothofagus forests, and are restricted to beech species Fig. 158
(Hudson 1934, Dumbleton 1932, Milligan 1974). In Length 1.8-8 mm Tarsal formula 4-4-4 or 5-5-5
contradiction to earlier beliefs, this species is a symptom
rather than the cause of beech mortality. Adults and larvae DIAGNOSIS. Body black, brown, or greyish, sometimes
of Maoraxia eremita occur in beech forests, but are not metallic green, broadly oval, short, stout, and usually
restricted to beech species. The larvae may be found in thin strongly convex dorsally; dorsal surface usually glabrous
dead branches of Nothofagus species (Milligan 1974), and and strongly glossy, rarely dull and pubescent with patches
also occur in elm trees, while adults have been collected by of paler hair and with short bristles. Head directed down-
beating scrub in summer (Hudson 1934, Dumbleton 1932, wards, more or less concealed from above. Antennae
Morgan 1966). 10- or 11-segmented, usually weakly clubbed, with
A single flying specimen of Buprestis aurulenta segments 3-7 gradually enlarged. Elytra often pointed
collected in Remuera, Auckland, represents a species posteriorly, often with distinct striae. Legs often retract-
possibly introduced from the north-west coast of the able into cavities of mesosternum, and prosternum with
U.S.A. The larvae bore in old wood of most of the a broad process fitting into cavity of mesosternum (Fig.
Californian pines, and are particularly attracted to fire 52, 53). Procoxae transverse, with cavities open. Tarsi
scars of exposed pitchy wood (Clark 1938). with segment 3 bearing an anteriorly directed mem-
branous appendage below.
RANGE. North ®, South ®, Stewart O
Kermadecs O, Three Kings O, Chathams O, Snares O SYNOPSIS. Watt (1982a) recorded 49 nominal native
Aucklands O, Campbell O, Antipodes O, Bounties O species of byrrhid from New Zealand and over 79, many
undescribed, in NZAC. Broun (1910b) recognised 8 gen-
EXAMPLE. BUPRESTINAE - Nascioides enysi (Sharp, era (including Nosodendron, now in the family Nosoden-
1877) (Fig. 157). dridae [33]); today there are 8 genera in two subfamilies -
BYRRHINAE and SYNCALYPTINAE. Aspects of the higher
REMARKS. Suggested colloquial name: jewel beetles. classification of Byrrhidae are discussed by Watt (1971).

-33-
 Adults have been collected from under stones and on Aucklands
Aucklands O, Campbell O, Antipodes O, Bounties O
Celmisia daisies in the mountains, and by beating moss-
covered trees or shrubs; larvae have been found on moss EXAMPLE. Parnida agrestis Broun, 1880 (Fig. 159).
and feeding on it (Broun 1910b, Hudson 1934). Kuschel
(1990) recorded 2 species at Lynfield, Auckland, collected REMARKS. Suggested colloquial name: hairy water
on logs and on fairly damp ground and stream beds with beetles.
moss and liverworts. In general adults and larvae occur on
mosses or liverworts, and some are found among the roots SELECTED REFERENCES. Brown (1981b, 1991a).
of higher plants (Lawrence & Britton 1991, 1994). They
may be collected on moss, on the ground, or on tree trunks
at night, as they appear to be nocturnal feeders. [22] Family ELMIDAE
Fig. 160
RANGE. North ®, South ®, Stewart ®
Kermadecs O, Three Kings ®, Chathams O, Snares ® Length 2.8-5.0 mm Tarsal formula 5-5-5
Aucklands ®, Campbell ®, Antipodes O, Bounties O DIAGNOSIS. Body narrowly oval, slightly pointed
posteriorly, black, greyish-black, or brownish-black,
EXAMPLE. BYRRHINAE - Liochoria huttoni Pascoe, sometimes slightly iridescent, moderately glossy or dull
1875 (Fig. 158). (when glossy often with a metallic sheen), with fine and
sparse pubescence, usually of a velvety appearance.
REMARKS. The New Zealand fauna is diverse, but is Head small. Antennae moderately elongate and
poorly understood at the generic and specific level, and the slender, sometimes appearing slightly bead-like. Eyes
group requires revision. prominent and protruding. Pronotum trapezoidal,
Suggested colloquial name: moss beetles. with sharp basal angles, narrower than elytral base.
Elytra with rows of punctures. Procoxae transverse,
SELECTED REFERENCES. Broun (1910b), Gressitt & their cavities open behind. Legs Iarge, stout. Tarsal
Samuelson (1964a), Watt (1971). segments simple, the last segment enlarged and with
moderately large claws.

[21] Family DRYOPIDAE SYNOPSIS. Watt (1982a) recorded 7 nominal native

Fig. 159 species of elmid, but some 16 species are now represented
in NZAC, many of them undescribed. All New Zealand
Length 3-5 mm Tarsal formula 5-5-5
elmids belong to a single genus of subfamily LARAINAE.
DIAGNOSIS. Body broadly oval, usually more or less Elmids are aquatic or semiaquatic, but none can swim.
cylindrical, bluntly triangular posteriorly, brown to Hudson (1934) reported adults of Hydora picea in river
black, glossy, covered with long, unruly, protruding beds running over stones at the water's edge, sometimes
hairs. Antennae short, with a comb-like asymmetrical taking wing with great rapidity. In general the adults may
club of 7 segments (Fig. 29). Pronotum with an be found on stones in river and stream margins, and the
impressed longitudinal groove on either side. Base of larvae live under stones near the water. Few larvae have
elytra with several impressions. Prosternal process been associated with adults, and careful rearing is needed
broadly and deeply received into mesosternum. to provide information for systematic studies.
Procoxae transverse, their cavities open. Legs large,
thick. Last tarsal segment enlarged and often as long as RANGE. North ®, South ®, Stewart ®
3 or all remaining segments, with large claws. Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
SYNOPSIS. Watt (1982a) recorded 4 native species of
dryopid in New Zealand, in 2 genera, whereas Hudson EXAMPLE. LARAINAE - Hydora picea Broun, 1880 (Fig.
(1934) had listed only 2 native species. 160).
Adults and larvae of Parnida agrestis may be found in
leaf litter, commonly in litter of hard beech (Nothofagus REMARKS. Our elmid fauna is relatively diverse but
truncata) forests. Adults of P. longulus occur under logs poorly known, and there are probably many undescribed
in very wet places, and those of P. agrestis in river beds, species. The southerrn Larainae are under revision by Dr
running over stones at the water's edge in hot sunshine Manfred Jäch (Naturhistorisches Museum, Vienna).
(Hudson 1934). Suggested colloquial name: riffle beetles.

RANGE. North ®, South ®, Stewart O SELECTED REFERENCES. Brown (1981 a, 1991 c),
Kermadecs O, Three Kings O, Chathams O, Snares O Spangler & Brown (1981).

-34--
34-
[23] Family LIMNICHIDAE SYNOPSIS. Heterocerids are represented in New Zea-
Fig. 161 land by a single native species in the type genus. The type
specimen is from Wedderburn in Central Otago, but the
Length 1-2.5 mm Tarsal formula 4-4-4 (Hyphalinae), species has also been found at Whatipu and Bethels Beach
5-5-5 (Limnichinae)
near Auckland, Ballantrae near Palmerston North, Hau-
DIAGNOSIS. Body minute to small, oval, convex, moana near Hastings, and in Northland.
moderately glossy, black, with short yellowish or greyish All known species live in tunnels in moist sand or mud,
pubescence often forming wavy patterns on pronotum and feed on algae and organic debris. Our species may be
and elytra. Frontoclypeal suture present; clypeus dis- found in mud alongside streams and rivers, where adults
tinct. Antennae slender, often with a 3-segmented club; may be collected by `shore washing' with water and col-
basalmost 2 segments enlarged; antennal insertion on lecting floating debris by net. Occasionally large numbers
base of frons. Pronotal base sinuate. Scutellum large. of adults have been caught in light traps near rivers.
Procoxae transverse, their cavities open behind. Meso-
sternum long and wide, well extended into procoxae; RANGE. North ®, South ®, Stewart O
metasternum with a distinct transverse suture. As in Kermadecs O, Three Kings O, Chathams O, Snares O
byrrhids, legs may be folded into ventral cavities. Aucklands O, Campbell O, Antipodes O, Bounties O

SYNOPSIS. Watt (1982a) reported 7 nominal native EXAMPLE. HETEROCERINAE - Heterocerus novaeselan-
species of limnichid and 1 adventive. The 2 limnichid diae Charpentier, 1968 (Fig. 162).
genera in New Zealand belong in subfamilies HYPHALINAE
(5 species) and LIMNICHINAE (3 species, with possibly a REMARKS. There are related species in Australia.
fourth undescribed). Suggested colloquial name: mud beetles.
Species of Hyphalus are known from the intertidal zone
on barnacle-encrusted rocks and beach rocks. Species of SELECTED REFERENCES. Charpentier (1968).
Limnichus occur under rocks at the margin of streams and
rivers, and in moist litter in forested areas (e.g., Notho-
fagus forest). [25] Family PTILODACTYLIDAE
Fig. 163
RANGE. North ®, South ®, Stewart O
Length 5-8 mm Tarsal formula 5-5-5
Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O DIAGNOSIS. Body oblong to elongate, pale brown
sometimes with a reddish tinge to dark brown, moderately
densely pubescent. Head small, strongly deflexed and
EXAMPLE. LIMNICHINAE - Limnichus nigripes Broun, concealed from above; frontal suture distinct. Antennae
1886 (Fig. 161). 11-segmented, filiform or serrate, moderately long
(Fig. 20). Eyes large and protruding. Pronotum small,
REMARKS. Very little is known about New Zealand's transverse, with a well developed basal locking
Limnichidae. mechanism consisting of a comb with smalI teeth or
Suggested colloquial name: lake beetles. crenulations; lateral carinae sharp and incomplete
anteriorly. Elytra striate. Scutellum heart-shaped, not-
SELECTED REFERENCES. Britton (1973, 1977), ched anteriorly. Procoxae conical, transverse, narrowly
Brown (1991b). separated, their cavities open behind. Tarsi simple;
tarsomere 4 small.
[24] Family HETEROCERIDAE SYNOPSIS. Watt (1982a) reported 2 nominal native
Fig. 162 species of ptilodactylid and a total of some 8 species,
several undescribed, represented in NZAC. As conceived
Length 3.8-4 mm Tarsal formula 4-4-4
here, the family includes 1 described species of Byrro-
DIAGNOSIS. Body robust, elongate, densely pubescent, cryptus (subfamily ANCHYTARSINAE) with 4-7 awaiting
the pubescence forming a variegated pattern. Head description, and possibly another undescribed genus
prognathous, with strong mandibles and with large (Kuschel 1990). Following the classification proposed by
labrum and clypeus. Antennae short and thick, with an Lawrence & Newton (1995), Brounia—hitherto placed in
elongate, apparently 7-segmented club. Pronotum Ptilodactylidae—is transferred to Chelonariidae [26].
strongly transverse. Elytra largely subparallel. Procoxae Adults are predacious, and occur on vegetation near
transverse, their cavities open behind. Tibiae flattened, water in wooded areas. Kuschel (1990) reported adults of
spinose. Byrrocryptus collected by Malaise trap near a stream,

— 35 —
underneath overhanging stream banks, and amongst fern RANGE. North ®, South O, Stewart O
and moss on a steep, seeping slope. Specimens of B. Kermadecs O, Three Kings O, Chathams O, Snares O
urquharti have been collected by beating vegetation near Aucklands O, Campbell O, Antipodes O, Bounties O
margins of streams. Ptilodactylid larvae from other
regions occur in wet habitats, and may be riparian or EXAMPLE. Brounia thoracica Sharp, 1878 (Fig. 164).
aquatic; they feed on rotting vegetation, including leaves,
roots, or dead wood (Lawrence 1991, LeSage & Harper REMARKS. According to Lawrence & Newton (1995)
1976). B. thoracica resembles some members of the Callirhipidae
more than it does species of Chelonarium or Pseudo-
RANGE. North ®, South ®, Stewart O chelonarium (Chelonariidae), but it shares the serrate
Kermadecs O, Three Kings O, Chathams O, Snares O tibiae, tarsal structures, wing venation, and type of male
Aucklands O, Campbell O, Antipodes O, Bounties O genitalia typical of chelonariids.
Suggested colloquial name: none.
EXAMPLE. ΑΝCΗΥΤΑRSIΝΑE — Byrrocryptus urquharti
Broun, 1893 (Fig. 163). SELECTED REFERENCES. Lawrence & Newton
(1995).
REMARKS. The New Zealand fauna of Ptilodactylidae
is probably richer than is apparent from its representation
in collections. The subfamily Anchytarsinae is almost Superfamily ELATEROIDEA
certainly paraphyletic, as suggested by Lawrence & The superfamily definition used here is after Lawrence &
Newton (1995). Byrrocryptus, which occurs also in Britton (1991, 1994) and Lawrence & Newton (1995).
Australia, was placed in this subfamily on the basis of a Adults of all elateroids lack a mandibular mola and a
larval synapomorphy—longitudinal division of expanded transverse metasternal suture, and have an unsophisticated
submentum into three parts—with four other genera wing-folding mechanism; larvae have a single stemma on
unknown in New Zealand (Lawrence & Newton 1995). either side of the head and a feeding mechanism adapted
The entire group is much in need of revision. for liquid diets (Lawrence & Britton 1991, 1994). These
Suggested colloquial name: comb-waist beetles. plant-feeders usually have the rear of the prosternum
elongated into a process that fits into the mesosternum
SELECTED REFERENCES. Stribling (1986). (Fig. 54). Phylogeny is discussed by Muona (1996).

[26] Family CHELONARIIDAE [27] Family EUCNEMIDAE

Fig. 164a,b [assisted by J. Muona]
Length 5-6 mm Tarsal formula 5-5-5 Fig. 165

DIAGNOSIS. Body narrowly oval, partly subparallel, Length 4-10 mm Tarsal formula 5-5-5
moderately convex, in lateral view with an anterodorsal
pronotal hump, black. Head small, strongly deflexed, DIAGNOSIS. Very similar to Elateridae [28], but dif-
concealed from above; frontal suture distinct. Anten- fering in having the labrum membranous and con-
nae 11-segmented, with basal 2 segments small and cealed beneath the clypeus. Body brown to black,
segments 4-11 pectinate, asymmetrical (Fig. 22). Eyes narrowly elongate, subparallel, pubescent, broadest at
large and protruding. Pronotum small, transverse, pronotum and tapering posteriorly. Front edge of pro-
with a welI developed basal locking mechanism con- sternum straight across and not lobed. Head deflexed.
sisting of a comb with small teeth; lateral carinae sharp Antennae 11-segmented, filiform or serrate. Pronotum
and complete. Elytra with faint indications of striae. with hind angles acutely produced posteriorly. Elytra
Scutellum small, oval. Procoxae transverse, their cav- striate. Clicking device present (consisting of prosternal
ities open behind; mesocoxae separated by a distance process fitting into mesosternal cavity), clicking ability
shorter than coxal width; metacoxae transverse, in contact well developed. Procoxae globular, their cavities open.
basally. Metasternal longitudinal suture present and Tarsomeres simple. First 5 ventrites connate.
transverse suture absent. Tarsi simple, with segment 3
bilobed beneath and segment 4 minute. SYNOPSIS. Watt (1982a) reported 20 nominal native
species of eucnemid from New Zealand. We estimate 22
SYNOPSIS. There is a single native species of chelon- species, including 2 undescribed, in 7 genera and 2 sub-
ariid, Brounia thoracica, in New Zealand. We have families – MELASINAE and MACRAULACINAE. Apart from the
examined only 1 specimen, collected in a Malaise trap in scattered original descriptions, very little information is
cattle-browsed forest at Hicks Bay, north of Te Araroa. available on the group.

-36-
 Adults may be found on forest growth, in leaf litter or well separated, the Iast 2 connected by a membranous
wood mould. Kuschel (1991a) reported 2 eucnemid suture.
species collected under bark of Pinus radiata logs lying in
the bush and in decayed wood in the Auckland area. Hud- SYNOPSIS. Watt (1982a) reported 132 native and 3
son (1934) recorded some species collected by beating adventive species of elaterid in New Zealand, belonging to
forest growth. May also be collected using interception 26 genera in 6 subfamilies — AGRYPNINAE, PITYOBIINAE,
traps, including Malaise traps, or by sweeping forest DENTICOLLINAE, LISSOMINAE (PROTELATERINAE), ELATERINAE,
vegetation. and CARDIOPHORINAE. For subfamily and tribal identification,
New Zealand larvae are known from dead wood of see Stibick (1979).
Metrosideros (rata). Most Australian larvae also occur in Adults occur on foliage of trees and shrubs, in rotting
dead wood (Lawrence & Britton 1991, 1994), and logs, under bark of dead trees, and on the ground, and may
Nearctic larvae of the group are known as cross-wood be collected by sweeping vegetation, collecting at light, or
borers because they cut characteristic mines across the netting in flight in the daytime. Kuschel (1990) collected
grain of wood (White 1983). adults at Lynfield, Auckland from vegetation by beating,
from Acacia and Eucalyptus, in bush litter and on low-
RANGE. North ®, South ®, Stewart ® growing plants at night, at light, in paddocks, in Malaise
Kermadecs O, Three Kings O, Chathams ®, Snares O traps, in leaf litter and prunings, under bark, from sifted
Aucklands O, Campbell O, Antipodes O, Bounties O decaying wood, and reared from abandoned boards of
Pinus radiata.
EXAMPLE. MELASINAE - Neocharis simplex Sharp, According to Lawrence & Britton (1991, 1994), in
1877 (Fig. 165). general there are three types of elaterid larvae: (a) sap-
rophagous — feeding on rotten wood; (b) phytophagous —
REMARKS. In early classifications the eucnemids were feeding mainly on roots of plants; and (c) predacious.
treated as a subfamily of Elateridae [28], but Broun treated Many phytophagous larvae, known as wireworms, are
them as a distinct family. Their phylogeny, classification, agricultural pests feeding on roots of cereals and other
biogeography, and biology (including New Zealand gen- plants. For example, Ctenicera larvae are soil-inhabiting
era) were recently reviewed by Muona (1991, 1993). wireworms, some of them damaging root vegetables such
Suggested colloquial name: false click beetles. as carrots. Thoramus larvae live in dead wood, including
standing dead trunks of Pinus radiata, where they prey on
SELECTED REFERENCES. Muona (1991, 1993), Sharp larvae of Cerambycidae. Most larvae are probably facul-
(1877b). tative predators on soil-inhabiting arthropods.

RANGE. North ®, South ®, Stewart

[28] Family ELATERIDAE Kermadecs ®, Three Kings ®, Chathams ®, Snares O
[assisted by J. Muona] Aucklands O, Campbell O, Antipodes O, Bounties O
Fig. 166, 167
Length 3-30 mm Tarsal formula 5-5-5 EXAMPLES. AGRYPNINAE - Thoramus wakefieldi Sharp,
1877 (Fig. 166); DENTICOLLINAE - Amychus candezei
DIAGNOSIS. Body elongate, narrow, cylindrical to Pascoe, 1876 (Fig. 167).
slightly flattened, usually brown or black, rarely spotted,
finely pubescent or nearly glabrous, often glossy, with REMARKS. This group is in need of taxonomic revision,
medial part usually subparallel and both ends rounded. as there is no comprehensive treatment available. The
Head with labrum sclerotised, visible and free, and general classification of Elateridae is in a state of flux, and
clypeus not distinct. Antennae 11-segmented, located the subfamilies are variously combined by different
near but not between eyes, usually sawtoothed or comb- authors (Lawrence & Newton 1995).
like, never clubbed. Prothorax loosely joining mesothorax Suggested colloquial name: click beetles (some larvae —
and capable of rapid movement on basal joint, which wireworms).
produces a jumping movement. Pronotum with hind
angles acutely produced posteriorly. Click mechanism SELECTED REFERENCES. Calder (1976, 1984, 1996),
always present (Fig. 54)—the most distinctive feature of Gaedike (1969), Sharp (1877b), Stibick (1979).
the family, consisting of prosternal process and mesoster-
nal cavity. Elytra usually subparallel for most of length,
tapering posteriorly. Procoxae globose, their cavity open [29] Family LYCIDAE
behind Tarsi simple, sometimes with setal brushes or
membranous appendages. Prosternum truncate or lobed in Fig. 168

front. Abdomen with 5 ventrites, rarely 6, all usually Length 9-10 mm Tarsal formula 5-5-5

—37—
DIAGNOSIS. Body black and orange, soft, flattened, distinctly shortened, exposing wings and many tergites,
loosely formed. Head small, partially concealed from often with inconspicuous ridges. Procoxae projecting,
above. Antennal bases in proximity; antennae thick, their cavities open. Legs long and slender. Tarsi with
strongly serrate, moderately long (Fig. 21). Eyes pro- segment 4 bilobed.
truding. Pronotum narrower at base than elytra, with
several ridges dividing disc into irregular depressions; SYNOPSIS. Watt (1982a) recorded 40 native and 1
base slightly sinuate. Elytra leathery, narrowly elongate, adventive species of cantharid in New Zealand, belonging
subparallel, expanded beyond body margins, with dis- to 3 genera in 2 subfamilies — DYSMORPHOCERINAE and
tinct round humeri and longitudinal ridges, between MALTHININAE. By far the most species (37) are placed in
ridges with a double row of large punctures. Procoxae Asilis (Wittmer 1979). A new subfamily classification was
projecting, their cavities open behind. Femora and tibiae proposed by Brancucci (1980).
slightly flattened. Tarsi with segment 3 and particularly Adults are diurnal and often seen on flowers and foliage
segment 4 bilobed. of various trees and shrubs, and occasionally may be found
under stones and bark. The majority of species are
SYNOPSIS. Watt (1982a) recorded a single species of believed to be predacious on other insects, but some feed
lycid from New Zealand, corresponding to Metriorrhyn- on plants, nectar, and pollen (LeSage 1991b). A common
chus erraticus Broun (1893a), described from Drury, method for collecting adults is sweeping and beating
Pokeno, and Clevedon. This is implicitly a synonym of vegetation.
Porrostoma rufipenne, an adventive Australian species, Kuschel (1990) reported 3 cantharid species from
reported by Kuschel (1990) from Lynfield, Auckland. Auckland, collected by sweeping grass and weeds, beating
Adults fly on warm days, and feed on flower nectar and vegetation, from flowers in a swamp, and in a Malaise trap.
pollen. Larvae may be found under the baτk of logs. Larvae are primarily nocturnal, preying chiefly on small
arthropods living in debris on the ground (LeSage 1991b).
RANGE. North ®, South ®, Stewart Ο In New Zealand some larvae have been extracted from leaf
Kermadecs O, Three Kings O, Chathams O, Snares O litter using Berlese funnels.
Aucklands O, Campbell O, Antipodes O, Bounties O
RANGE. North ®, South ®, Stewart O
EXAMPLE. Porrostoma rufipenne (Fabricius, 1801) Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties Ο
(Fig. 168).
EXAMPLE. DYSMORPHOCERINAE - Asilis fulvithorax
REMARKS. The bright colours of, e.g., Porrostoma rufi- (Broun, 1880) (Fig. 169).
penne may be a warning signal to potential predators, since
lycids in general are believed to mimic the colours of REMARKS. Suggested colloquial name: soldier beetles.
insects distasteful to predators.
Suggested colloquial name: net-winged beetles. SELECTED REFERENCES. Brancucci (1980), LeSage
(1991b), Wittmer (1979).
SELECTED REFERENCES. Βοcák & Βocáková (1990).

Series BOSTRICHIFORMIA
[30] Family CANTHARIDAE We have followed the concept of Lawrence & Britton
Fig. 169 (1991, 1994) and Lawrence & Newton (1995) in defining
Length 1.9-7 mm Tarsal formula 5-5-5 this group. It is considered to be a paraphyletic assemblage
consisting of Polyphaga with strong affiliation to the
DIAGNOSIS. Body usually uniformly black or brown, cucujiform lineage, but lacking shared apomorphies with
sometimes with pronotum yellowish to orange, soft, the series Cucujiformia. Jacobsoniidae are tentatively
narrowly oval to subparallel, slightly convex to flattened, included here because they lack some important syn-
moderately densely pubescent, often slightly glossy. Head apomorphies of Cucujiformia (Lawrence & Newton
large, deflexed but partially visible from above. Eyes 1995).
protruding. Labrum membranous and inconspicuous.
Antennae filiform (Fig. 17) or serrate, with insertions
welI separated. Pronotum transverse, broadly oval or Superfamily DERODONTOIDEA
approximately subquadrate, sometimes with 2 small, Members of this group are distinct in having paired ocelli
round basal impressions, and sometimes emarginate. on the frons, an elaborate system of depressions and
Elytra narrowly oval, often subparallel for most of usually ridges and canals on head and pronotum, and
length, usually loosely covering abdomen but sometimes elytral striae with deep basal pits.

-38-
[31] Family DERODONTIDAE (1991, 1994) and Lawrence & Newton (1995) considered
Fig. 170 their placement of Jacobsoniidae in Bostrichoidea to be a
tentative arrangement, as at present there is only weak
Length 2.0-2.4 mm Tarsal formula 5-5-5
evidence of some similarity between the two groups in
DIAGNOSIS. Body small, approximately oval in out- aedeagal characters.
line, brown or black, moderately glossy, with incon-
spicuous short adherent pubescence. Head with paired
ocelli and usually elaborate depressions. Antennae 11- [32] Family JACOBSONIIDAE
segmented, the basalmost 2 segments enlarged, and Fig. 171
with a 3-segmented club. Eyes protruding, coarsely
Length 0.7-2 mm Tarsal formula 5-5-5 or
faceted. Pronotum distinctly narrower than base of elytra, 3-3-3 (appearing 2-2-2)
with a series of depressions and usually with ridges
dividing disc into several sections. Elytra broadly oval, DIAGNOSIS. Body small to minute, elongate, reddish-
with deeply impressed striae each usually having a deep pit brown or brown, glossy, usually punctate, with (unique
at base. Procoxae projecting, their cavities usually character) metasternum at least as long as 5 visible
closed behind; metacoxae transverse, with well devel- sternites combined, giving the appearance of hind legs
oped plates. emerging near the end of the abdomen. Antennae with a
1-3-segmented club. Procoxae projecting, their cav-
SYNOPSIS. There are 2 native species of derodontid in ities usually open.
New Zealand, both in genus Nothoderodontus of subfam- Of the two distinct groups in our fauna the first,
ily LARICOBIINAE (Watt 1982a, Lawrence 1985, Lawrence represented by a single species in genus Saphophagus, has
& Newton 1995). the body larger (length 1.9-2 mm) and tarsi 5-5-5-seg-
Nothoderodontus gourlayi feeds on hyphae and conidia mented. The second comprises possibly three undes-
of sooty moulds (Ascomycetes: Capnodiaceae and related cribed native or introduced species in genus Derolathrus,
families) (Lawrence 1985). Specimens including larvae distinct by their minute body size (length 0.7-0.8 mm),
have been taken in Nothofagus forest from sooty mould yellowish rust-brown colour, antennae appearing to have a
growing on the honeydew excreted by a scale insect. 1-segmented club, head abruptly constricted behind
eyes to form short temples, oval pronotum, and narrowly
RANGE. North O, South ®, Stewart O oval elytra, tapering distally.
Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O SYNOPSIS. Watt (1982a) recorded 1 described native
species and 2 other species of jacobsoniid housed in
EXAMPLE. LARICOBIINAE - Nothoderodontus gourlayi NZAC. These are grouped in 2 genera — Saphophagus,
Crowson, 1959 (Fig. 170). with 1 native species known only from the South Island,
and Derolathrus, with 1 undescribed species (or possibly
REMARKS. Derodontids are considered to be among the 3), collected in Omahuta State Forest, Northland, and on
most primitive of Polyphaga (Lawrence & Newton 1995). Codfish Island and Rurima Island in the far south. Dero-
Suggested colloquial name: sooty fungus beetles. lathrines are also known from Norfolk Island, Lord Howe
Island, Christmas Island, and Australia (Lawrence &
SELECTED REFERENCES. Crowson (1959), Lawrence Britton 1991, 1994, Löbl & Burckhardt 1988). Crowson
(1985), Lawrence & Hlavac (1979). (1959) considers S. minutus to be one of the most remark-
able of New Zealand's endemic insects, possibly a relict
coeval with the tuatara, Sphenodon.
Superfamily BOSTRICHOIDEA Adults of S. minutus occur under loose bark of dead
The concept used here is after Lawrence & Britton (1991, Nothofagus branches. Larvae of S. minutus described by
1994) and Lawrence & Newton (1995). The superfamily Crowson (1959) were found under moist and close-fitting
comprises five very diverse families (Anobiidae, Bostri- bark of dead Nothofagus or, in one instance, of pukatea
chidae, Dermestidae, Jacobsoniidae, and Nosodendridae), (Laurelia novae-zelandiae). Specimens of Derolathrus
all with 5-5-5-segmented tarsi. Lawrence & Newton were found in guano and in a nest of kiore, Rattus exulans.
(1982) and Lawrence & Britton (1991, 1994) proposed
inclusion of the Dermestidae in Bostrichoidea on the basis RANGE. North ®, South ®, Stewart ®
of modified cryptonephridism, similar aedeagal structures, Kermadecs O, Three Kings O, Chathams O, Snares O
and lack of a basal mandibular mola in larvae. Ivie (1985) Aucklands O, Campbell O, Antipodes O, Bounties O
added Nosodendridae to this superfamily on the grounds
of assumed secondary loss of cryptonephridism in both EXAMPLE. Saphophagus minutus Sharp, 1886 (Fig.
Dermestidae and Nosodendridae. Lawrence & Britton 171).

-39-
REMARKS. The taxonomic position of this small family [34]
[34] Family DERMESTIDAE
is uncertain (Lawrence & Newton 1995). Fig. 173
Suggested colloquial name: Jacobson's beetles.
Length 3-10 mm Tarsal formula 5-5-5
SELECTED REFERENCES. Crowson (1959), Löbl & DIAGNOSIS. Body compact, usually robust, oblong,
Burckhard (1988). elliptical, or broadly ovate, usually brown to black, rarely
reddish-brown (Trichelodes), sometimes with pale spots,
slightly to strongly glossy, clothed with short or long,
[33] Family NOSODENDRIDAE erect or decumbent hairs or scales, these often forming
Fig. 172 patterns. Head deflexed and more or less concealed
from above, usually with a median ocellus. Antennae
Length 4-7 mm Tarsal formula 5-5-5
filiform or pectinate, short and clubbed, the club
DIAGNOSIS. Body black, ovoid, strongly convex, usually 3-segmented but sometimes 4-8-segmented;
glabrous or with sparsely distributed short bristles, antennae often fitting into grooves below either side of
moderately to strongly glossy. Head prognathous and pronotum. Eyes moderate-sized to large and protruding.
not retractable; clypeus narrow but visible; maxillae and Pronotum transverse, narrower than elytral base,
prementum concealed beneath enlarged mentum. Antennae sometimes strongly convex medio-apically, giving a
11-segmented, with a 3-segmented tomentose club. hump-backed appearance laterally, usually with sharp
Pronotum strongly transverse (Fig. 118). Scutellum tri- lateral margins (absent in Trichelodini). Procoxae trans-
angular, large. Elytra broadly curved laterally, rounded verse, conical or globose and usually projecting, their
posteriorly, with punctate or setose striae. Prosternum cavities open; metacoxae usually excavated for
with a triangular process fitting into mesosternal reception of femora (Fig. 56). Tarsal segments simple.
cavity. Procoxae transverse, their cavities open. Legs
capable of folding into ventral cavities. Tarsal segments SYNOPSIS. Watt (1982a) reported 11 native and 6
simple. adventive species of dermestid in New Zealand. They
belong in 7 genera representing 4 subfamilies — DER-
SYNOPSIS. Watt (1982a) recorded 2 native species of MESTINAE, TRINODINAE, ATTAGENINAE, and MEGATOMINAE.
nosodendrid in New Zealand, whereas Hudson (1934) and Species of Anthrenocerus, Anthrenus, Attagenus, Der-
Endrödy-Younga (1989) reported 3 species. There are 2 mestes, and Trogoderma especially are economically
species represented in NZAC, Nosodendron zealandicum important, and are represented here mostly by adventive
and the smaller N. ovatum. cosmopolitan species. According to Crowson (1959)
Adults are encountered infrequently in slime fluxes on native New Zealand species represent only 2 of the 6
wounded trees, where they probably feed on bacteria and subfamilies that he recognised, Trinodinae (Trichelodes)
products of fermentation. Specimens of N. ovatum have and Anthreninae (Trogoderma). Kuschel (1990) recorded
been collected from under rotten branches, from litter, and 8 species from the Auckland area, most with adults taken
one from bat guano; two specimens of N. zealandicum from flowers.
were taken in a light trap. No habitat data for our larvae are In general adult dermestids usually feed on nectar or
available, but Lawrence & Britton (1991, 1994) reported pollen (e.g., Anthrenus, Trogoderma), though some spe-
larvae of Australian species living in slime fluxes. cies are known to use the larval food sources or do not take
food at all (Booth et al. 1990, Lawrence & Britton 1991,
1994). Larvae are primarily scavengers, feeding on dried
RANGE. North ®. South O, Stewart O animal remains such as bones, meat or carrion, hides, furs,
Kermadecs O, Three Kings ®. Chathams O, Snares O leather, wool and woollen products, silk, feathers, dried
Aucklands O, Campbell O, Antipodes O, Bounties O insects, spider webbing, nests of birds and mammals, and
even wasps' nests (Reesa vespulae — Waller & Watt 1979),
EXAMPLE. Nosodendron zealandicum Sharp, 1882
and some are known from stored grain. Some species of
(Fig. 172).
Trogoderma can subsist partly or entirely on vegetable
matter. Larvae of Trogoderma were reported by Crowson
REMARKS. Adults of Nosodendridae may be readily (1959) from a number of unspecified New Zealand
distinguished from the externally similar Byrrhidae [20] localities, in cavities and under dry bark, mainly of
by their prognathous (not retractable) head, distinct cly- Podocarpaceae, and especially rimu (Dacrydium cupres-
peus, and enlarged mentum. sinum).
Suggested colloquial name: wounded tree beetles. For cosmopolitan pest species of this group see Booth
et al. (1990), Bousquet (1990), Harney (1993), and Law-
SELECTED REFERENCES. Crowson (1959), Endrödy- rence & Britton (1994). Adventive species in New
Younga (1989). Zealand include the carpet beetle, Anthrenus verbasci,

— 40--
40—
larvae of which can seriously damage untreated woollen The third type (Fig. 176), represented by members of
carpets and garments; adults may be found on flowers in the LYCTINAE, is distinguished as follows. Body elongate
spring. The Australian carpet beetle, Anthrenocerus aus- but flattened. Head prognathous, visible from above.
tralis, has rather similar habits. The hide beetle, Dermestes Antennae with a compact 2-segmented club. Pronotum
maculatus, can seriously damage hides and skins. When approximately trapezoidal, broadest apically, with
ready to pupate, its larvae bore into wood or other dense distinct lateral carinae. Elytra subparallel, with par-
material, often causing damage to structural timbers. allel rows of hairs. Tarsi with basalmost 4 segments
equally short.
RANGE. North ®, South ®, Stewart O
Kermadecs ®, Three Kings O, Chathams ®, Snares O SYNOPSIS. Watt (1982a) reported 1 native, 2 adventive,
Aucklands O, Campbell O, Antipodes O, Bounties O and 1 undescribed species of bostrichid (including Lyc-
tinae) in New Zealand. Our current estimate indicates 8
EXAMPLE. MEGATOMINAE - Trogoderma maestum species, in 4 genera and 3 subfamilies — DINODERINAE,
Broun, 1880 (Fig. 173). LYCTINAE, and EUDERINAE (see Remarks for details).
Euderia is represented by a single native species, and the
REMARKS. An economically important group with remaining genera by adventive pest species.
several serious pests of hides, wool products, museum Adults of Euderia squamosa have been collected by
collections, and stored grain products. The dry exuviae beating Nothofagus branches, and larvae have been found
and spear-headed hairs of the larvae can cause asthma or in galleries in moist or sappy bark of dead or injured trees.
allergies. A detailed description of the bostrichid larva is provided
Suggested colloquial name: hide beetles. by Crowson (1961). Elsewhere, bostrichids are known to
bore into sapwood and bamboo.
SELECTED REFERENCES. Beal (1991), Crowson
(1959), Mroczkowski (1968), Peacock (1978, 1993). RANGE. North ®, South ®, Stewart O
Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
[35] Family BOSTRICHIDAE
(incl. LYCTINAE) EXAMPLES. EUDERINAE - Euderia squamosa Broun,
Fig. 174-176 1880 (Fig. 174); DINODERINAE - Dinoderus minutus
(Fabricius, 1775) (Fig. 175); LYCTINAE — Lyctus brunneus
Length 3-7 mm Tarsal formula 5-5-5 (Stephens, 1829) (Fig. 176).
DIAGNOSIS. There are three distinct morphological
types of bostrichid in our fauna. REMARKS. There are several unreported, probably
The first type (Fig. 174), represented by the monotypic recently adventive species of Lyctinae and Bostrichinae in
genus Euderia (EUDERINAE), may be characterised as NZAC which might become of economic importance to
follows. Body (except pronotum) elongate and slightly the timber industry. Present holdings are: 1 species of
flattened; dorsal surface with protuberances and Euderiinae, Euderia squamosa (Fig. 174); 3 species of
depressions, and covered with white and brown scales Dinoderinae — Dinoderus minutus (pantropical pest of
forming a pattern. Antennae with a 3-segmented club, bamboo and maize) (Fig. 175), Rhizopertha dominica (a
consisting in males of long flabellate branches and in major economic pest in a variety of dried food products),
females of shorter asymmetrical segments. Pronotum and one undetermined; and possibly 4 species of Lyctinae
narrow, irregularly cylindrical, as broad as head. Eyes —Lyctus brunneus, a cosmopolitan pest in a wide variety of
large, protruding. Elytra subparallel, nearly twice as timbers (Fig. 176), L. planicollis, L. linearis, and one of
broad at base as pronotum, and with distinct shoulders. uncertain identity.
Tarsi with basal segment as long as the following two One of the most interesting bostrichid species in our
combined. fauna is Euderia squamosa. According to Lesne (1934,
The second type (Fig. 175) is represented by mainly 1938) it represents the most primitive surviving bostrichid
introduced species of DINODERINAE (Dinoderus, Rhizo- type. Crowson (1961) considers Euderia to be more
pertha), and may be characterised as follows. Body closely related to Bostrichinae than to any other group, and
cylindrical, similar to Scolytinae [82]. Head partially or to be regarded as a probable relict of a group from which
completely concealed from above by the hood-like the Bostrichinae were derived. Euderia squamosa has also
extended pronotum. Antennae with a 3-segmented some anobiid features (Crowson 1961), and was originally
asymmetrical club. Pronotum with rasplike teeth at placed by Broun (1880) in the Anobiidae [36].
front. Elytra often obliquely cut posteriorly, and For a key to subfamilies of bostrichid adults, see Crow-
usually bearing spines or lateral teeth. son (1961).

—41—
 Suggested colloquial name: twig borer beetles (Lyc- Adults and larvae may be found in dead trees, logs, or
tinae — powder post beetles). timber, though some species bore into furniture, organic
stored products, fungal fruiting bodies, and dried organic
SELECTED REFERENCES. Bain (1978), Crowson material of animal and plant origin. Larvae are generally
(1961), Lesne (1934). wood and bark borers, but larvae of Dorcatominae feed on
woody or fibrous fungi. Larvae of Ptininae are generally
found in dry organic material of animal or plant origin,
[36] Family ANOBIIDAE e.g., leaf litter, and some Australian species are known to
(incl. PTININAE) occur in nests of birds and social insects (Lawrence &
Fig. 177, 178 Britton 1991, 1994). Some ptinines are widespread, cos-
Length 3-9 mm Tarsal formula 5-5-5 mopolitan species recognised as minor pests of stored
products, woollen goods, and book bindings (Booth et al.
Of at least 3 distinct subfamilies in New Zealand, the 1990, Bousquet 1990, Harney 1993). One of the adventive
Ptininae are readily distinguishable by their spider-like European ptinines in New Zealand, Ptinus tectus, infests
appearance. Ptininae also have approximated antennal stored foods. Hudson (1934) reported adults of some
insertions, lack lateral pronotal carinae, and have species collected by beating forest growth, amongst kiekie
widely separated metacoxae. In contrast most Ano- (Freycinetia), and from dry timber of houses (Anobium
biinae have a hooded prothorax, usually completely ruficorne, Stegobium paniceum). In New Zealand Ano-
concealing head from above, and contiguous metacoxae. bium punctatum may cause serious damage to untreated
structural timbers, and occasionally to furniture. The
DIAGNOSIS. Body pale brown, dark brown, or grey- several species recorded in the Auckland area by Kuschel
ish, usually elongate and cylindrical, sometimes ovate (1990) were collected from wallboards of old homes and
(Anobiinae, Dorcatominae) or. spider-like and often sheds, wood and foliage of native and introduced trees and
multicoloured (Ptininae), clothed with decumbent or shrubs, and leaf litter.
erect hairs or with scales, these often forming colour
patterns (Ptininae), rarely appearing glabrous. Head de- RANGE. North O, South O, Stewart O
flexed and usually concealed from above (Fig. 10). Kermadecs O, Three Kings O, Chathams ®. Snares O
Antennae 9-11-segmented, filiform and without a Aucklands O, Campbell O, Antipodes ®. Bounties O
distinct club (Ptininae), or usually serrate or pectinate,
with a 3-segmented club, its segments asymmetrical, EXAMPLES. ANOBIINAE - Hadrobregmus magnus
lengthened, and expanded. Prosternum reduced, often (Dumbleton, 1941) (Fig. 177); PTININAE - Ptinus speci-
deeply excavate, with cavity continuing on mesosternum. osus Broun, 1880 (Fig. 178).
Procoxae globose or projecting; metacoxae usually
contiguous, excavated to receive femora and with coxal REMARKS. According to Ivie (1985) the Anobiidae
plates in Anobiinae, not excavated and without coxal should be considered as a subfamily of Bostrichidae [35].
plates in Ptininae. Abdomen with usually 5 ventrites. For information on the species of economic importance
see Hinton (1941), Booth et al. (1990), Bousquet (1990),
SYNOPSIS. The world fauna of anobiids (in the sense of and Harney (1993).
Lawrence & Britton 1991, 1994) comprises some 180 Suggested colloquial names: borer beetles, furniture
genera with over 2000 species; their higher classification beetles (Ptininae — spider beetles).
requires revision. A modern subfamily arrangement is
provided by Lawrence & Newton (1995), and a key to the SELECTED REFERENCES. Español (1976a, b, 1979),
subfamilies of Anobiidae of the world (older concept, Hosking (1976, 1978f), Milligan (1977, 1979a), White
excluding Ptinidae) is provided by White (1974). (1974).
In New Zealand there are 23 native and 5 adventive
nominal species of ANOBIINAE and DORCATOMINAE (ori-
ginally in Anobiidae), and 5 native and 6 adventive species Series CUCUJIFORMIA
of PTININAE (originally in Ptinidae) (Watt 1982a); these We have followed the concept of Lawrence & Britton
belong to some 25 genera. Kuschel (1990) recorded 9 (1991, 1994) and Lawrence & Newton (1995) in defining
species of Anobiidae including 1 ptinine from Lynfield, this series. Adults are characterised by acone eyes, crypto-
Auckland. Five species in 2 genera have been recorded nephridial Malpighian tubules, lack of functional spiracles
from the Kermadec Islands, with some extralimital records on abdominal segment 8, metacoxae not excavate, absence
from Norfolk Island and Lord Howe Island (Español of metacoxal plates, reduction of pregenital segments 9
1979). Ptinus tectus has been recorded from the Auckland and 10, and the form of the male genitalia. Larvae have
Islands, Antipodes Islands, Campbell Island, and The many primitive polyphagan features; usually the galea and
Snares (Watt 1962, 1971). lacinia are fused into a single mala.

-42-
Superfamily CLEROIDEA 24 species in 10 genera is endemic except for the adventive
Characterised by adults with dorsum of body usually stored products pest Tenebroides mauritanicus.
covered with protruding bristly hairs, projecting procoxae, Most species of Trogossitidae are predacious as larvae
and 5-segmented tarsi with segments often lobed, and and adults (Lawrence & Britton 1991, 1994), but probably
larvae with a single pretarsal seta, lack of mandibular mola all (or at least some) adults of the tribe Rentoniini are not.
in all stages, and usually predacious habit (Lawrence & These usually occur in leaf litter of native forests, under
Britton 1991, 1994, Lawrence & Newton 1995). Larvae loose bark, or on flowers of Pinus radiata (one record); at
usually have a basal mandibular process and a pedunculate least some of them feed on pollen (Crowson 1966a). The
seta on the mala (Lawrence & Newton 1995). single known rentoniine larva is suspected to occupy an
A key to the families of Cleroidea (adults and larvae) is arboreal habitat (Crowson 1966a). The remaining species
provided by Crowson (1964). will probably be most common under bark of dead trees.
Larvae of many species of Lepidopteryx (formerly as
Leperina) are predators in this habitat, and occur in
[37] Family TROGOSSITIDAE association with the adults.
(Incl. PELTIDAE) Collecting: sifting leaf litter and processing it through
Fig. 179-181 Berlese funnels, beating forest foliage, or picking adults
from under bark. Kuschel (1990) collected eight species of
Length 0.6-18 mm Tarsal formula 5-5-5 Rentoniinae in Auckland, from vegetation, decayed wood,
DIAGNOSIS. We have followed the classification pro- ground litter, and stream-bed moss.
posed by Lawrence & Newton (1995).
General characteristics: antennae 11-segmented, RANGE. North ®, South ®, Stewart
with a loose 3-segmented club sometimes appearing Kermadecs O, Three Kings ®, Chathams ®, Snares O
asymmetrical; procoxal cavities open or closed; ab- Aucklands O, Campbell O, Antipodes O, Bounties O
domen with 5 visible segments. Three main body forms
may be recognised in our trogossitid fauna, as follows. EXAMPLES. RΕΝΤΟΝΙΙΝΑE - Rentonidium costiventris
Crowson, 1966 (Fig. 179); LOPHOCATER1NAE - Grynoma
PROTOPELTINAE: body medium-sized (length 2.6-4
regularis Broun, 1886 (Fig. Ι 80); TROGOSSITINAE -
mm), of nitidulid form, broadly oval, subparallel Lepidopteryx nigrosparsa (White, 1846) (Fig. 181).
medially, covered with long hairs; antennae with a
symmetrical, oval, 3-segmented club; elytra strongly REMARKS. The higher classification of Trogossitidae is
punctate but without striae. not settled. An important step in classification of the group
RENTONIINAE (Fig. 179): small to minute beetles was the establishment by Crowson (1964, 1966a) of the
(length 0.6-1 mm), superficially similar to clambids but trogossitid complex within the superfamily Cleroidea.
readily distinguishable by the lack of metacoxal plates; Crowson (1966x) recognised family Peltidae as having 4
body oval to nearly spherical, strongly convex and subfamilies, of which Rentoniinae was erected for 5 New
glossy, glabrous or with short and sparse pubescence; Zealand genera. Subsequently Crowson (1970) divided
colour brown to black or sometimes pale reddish brown; the trogossitids (in a broad sense) into 3 distinct families,
head broad and deflexed; pronotum broadly rounded distinguishing the Peltidae and Trogossitidae by differ-
anteriorly, with front angles not produced; scutellum ences in their mouthparts, the latter group having adult
moderately large. mandibles without a definite molar part and the lacinia
LOPHOCATERINAE and TROGOSSITINAE (Fig. 180, 181): without an apical hook, and larval mandibles with a
smalI to medium-sized beetles (length 3-18 mm) different lacinia mandibulae' and the maxillary mala with
superficially similar to tenebrionids, but readily a pedunculate seta. The classification of Trogossitidae and
distinguishable by 5-5-5-segmented tarsi and other relationships of the higher taxa of Cleroidea are discussed
characters; body broadly to narrowly oval; elytra often by Barron (1971) and Slipinski (1992). Lawrence &
subparallel, flattened to moderately convex; upper Britton (1991, 1994) merged the 3 trogossitid families of
surface with short or long hairs, bristles, or scales, Crowson (1970) into the single family Trogossitidae.
often bicoloured and spotted; colour brown to black with Suggested colloquial name: cadelle beetles (peltines -
paler spots (greyish, yellowish, etc.), moderately glossy to shield beetles).
dull; head moderately narrow and prognathous; pro-
notum usually with anteriorly produced front angles. KEY TO SUBFAMILIES OF TROGOSSITIDAE
OCCURRING IN NEW ZEALAND
SYNOPSIS. Lawrence & Newton (1995) recognise trog- [Modified from Slipinski 1992]
ossitids as comprising 9 subfamilies, of which 4 are
represented in New Zealand - PROTOPELTINAE, RENTONIINAE, 1 Procoxal cavities externally closed
LOPHOCATERINAE, and TROGOSSITINAE. Our fauna of some ... (Fig. 181) .. TROGOSSITINAE

—43—

—Procoxal cavities externally open Suggested colloquial name: southern cadelle beetles.

2(1) Protibia with 1 spur; antennal club markedly asym- SELECTED REFERENCES. Watt (1979b).
metrical ...(Fig. 180) .. LOPHOCATERINAE
—Protibia with 2 unequal spurs; antennal club sym-
metrical 3
...
[39] Family CLERIDAE
Fig. 183, 184
3(2) Very convex and rounded, clambid-like; procoxal
cavities internally closed (Fig. 179) .. RENTONIINAE
...
Length 5-20 mm Tarsal formula 5-5-5,
—Never very convex and rounded, not clambid-like; appearing 4-4-4
procoxal cavities internally open PROTOPELTINAE
...
DIAGNOSIS. Body elongate, usually subparallel, often
subcylindrical, with dorsal surface usually coarsely
SELECTED REFERENCES. Crowson (1964, 1966a, punctate. Coloration dark to pale brown, black, violet,
1970), Foster (1991a), Slipinski (1992). yellow, often spotted with yellow or pale brown, rarely
multicoloured, often with a metallic tinge. Setation of
variable length, short and decumbent or long and erect,
[38] Family CHAETOSOMATIDAE uniformly brown or yellow, and often forming a colour
Fig. 182 pattern. Head slightly deflexed, usually insignificantly
broader than pronotum. Antennae short, sawtoothed or
Length 7-14 mm Tarsal formula 5-5-5
filiform, most often with a 3-segmented club consisting
DIAGNOSIS. Body rectangulate, at least 4x longer of elongate and often asym-metrical segments. Eyes large,
than broad, subdepressed, coarsely punctate, glossy, often protruding, slightly to strongly emarginate.
with exceptionally long and protruding hairs (often Labial palps with terminal segment often enlarged. Pro-
longer than width of elytron). Head strongly prognathous, notum usually narrower than elytral base (exception:
with pronounced mandibles. Antennae 11-segmented, Necrobia), broadest medially and of equal width basally
uniform in width, about as long as forebody, the and apically, usually without lateral carinae (with carinae
segments bead-shaped. Eyes small but slightly protru- in Metaxina). Elytra subparallel, pear -shaped or grad-
ding. Pronotum subquadrate, in our fauna without setal ually broadening apically (exception: abbreviated in
tufts; front angles pointing slightly anteriorly. Elytra Paupris), irregularly and more or less coarsely punctate.
subparallel and elongate, with humeral angles acute Procoxae projecting, their cavities open or closed. Legs
(Chaetosoma) or obtuse (Chaetosomodes), and with long- long. Mesotrochanters oblique (Fig. 58). Tarsi usually
itudinal rows of punctures. Scutellum oblong. Procoxae with 1 or 2 segments lobed (simple in Metaxina); basal
globose and slightly transverse, their cavities open. segments sometimes reduced. Abdomen with 5 or 6
Legs with distinct trochanters. Tarsal segments simple. ventrites.
Pygidium slightly exposed.
SYNOPSIS. Watt (1982a) reported 37 native and 2
SYNOPSIS. Chaetosomatids are known to occur in New adventive species of clerid in New Zealand. These rep-
Zealand and Madagascar (Ekis & Menier 1980). There are resent some 8 genera in 6 subfamilies – THANOCLERINAE,
2 endemic genera in New Zealand, Chaetosoma with 1 PHYLLOBAENINAE, CLERINAE, ENOPLIINAE, TARSOSTENINAE,
described species and 2 undescribed (NZAC) and Chaeto- and KORYNETINAE. Phymatophaea is the largest New
somodes with 1 described species. Zealand genus, containing the majority of species. Our
Adults and larvae may be found under bark of dead endemic genus Metaxina is the most divergent, and there is
trees, logs, etc., where they feed on other wood-boring some confusion as to its phylogenetic affiliations; it
insects. The larva of Chaetosoma scaritides inhabits bur- probably represents a primitive lineage, and may be
rows in Nothofagus and other trees. considered a relict in the New Zealand fauna (Crowson
1964). Kuschel (1990) collected 5 clerid species in Auck-
RANGE. North ®, South ®, Stewart O land, including the 2 adventive ones.
Kermadecs O, Three Kings O, Chathams ®, Snares O Adults of many New Zealand native species are
Aucklands O, Campbell O, Antipodes O, Bounties O commonly found on flowers and leaves in sunny locations,
where they prey on other insects. The predominant habitat
EXAMPLE. Chaetosoma scaritides Westwood, 1851 of larvae is dead wood, and their food consists of larvae of
(Fig. 182). woodborers such as Anobiidae, Bostrichidae, Buprestidae,
Cerambycidae, or Scolytinae (Crowson 1964). The cos-
REMARKS. Their present widely disjunct distribution mopolitan species Necrobia ruficollis and N. rufipes occur
suggests that chaetosomatids are an old group of Gond- on carrion and stored products of animal origin, and have
wana origin. been reported as feeding on a cargo of copra (Watt 1975).

–44–
 Collecting: netting adults in flight, sweeping vegetation REMARKS. For a key to the adults of all described
particularly when in flower, and sifting bark and forest species of Phycosecis, see Crowson (1964).
debris for larvae. Suggested colloquial name: beach beetles.

RANGE. North ®, South ®, Stewart SELECTED REFERENCES. Crowson (1964), Watt
Kermadecs ®, Three Kings ®, Chathams ®, Snares O (1975, 1983a).
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLES. THANEROCLERINAE - Metaxina ornata Broun, [41] Family MELYRIDAE
1909 (Fig. 183); ΕNOΡLIINΑE - Phymatophaea violacea (also as DASYTIDAE)
(Fabricius, 1775) (Fig. 184). Fig. 186
REMARKS. A generic key to adults and larvae of New Length 2.5-6 mm Tarsal formula 5-5-5
Zealand taxa is provided by Crowson (1964), with a key DIAGNOSIS. Body soft, oblong to narrowly elongate,
to the subfamilies of adult Cleridae. For cosmopolitan dorsally flattened, usually with decumbent hairs and
species of economic importance, see Booth et al. (1990). often with additional scattered erect setae, uniformly
Suggested colloquial name: checkered beetles. dark-coloured in black, grey, brown, greenish, or blue,
often with a metallic tinge, usually glossy; dorsal surface
SELECTED REFERENCES. Crowson (1964), Foster of forebody sometimes with meshed microsculpture.
(1991b). Head often depressed medially, not or scarcely rost-
rate, usually comparable in size to pronotum. Labrum
well sclerotised; segments of maxillary palp not enlarged.
[40] Family PHYCOSECIDAE Antennae usually filiform, without a club. Eyes entire,
Fig. 185 moderately large, slightly protruding. Pronotum dis-
Length 2.5-3 mm Tarsal formula 5-5-5 tinctly narrower than elytral base (slightly broader than
width of elytron), often broadest at middle, often with
DIAGNOSIS. Body elongate oval, convex, variable in depressions. Elytra entirely covering abdomen or
colour, mostly dulI black, but with most of pronotum slightly shorter, exposing abdominal apex, subparallel
and head bearing whitish scales. Head prognathous, and rounded posteriorly or gradually broadening towards
partially concealed by semicircular anterior margin of apex. Procoxae projecting, their cavities open. Legs
pronotum, with 2 angular projections between eye and long and slender. Tarsal segments simple; terminal
base of antenna. Antennae appearing 10-segmented, segments bearing claws usually with membranous
with a 1-segmented club (club actually appears 2- appendages. Abdomen with 6 ventrites.
segmented). Pronotum horseshoe-shaped in outline,
with front angles strongly produced anteriorly. Elytra SYNOPSIS. Watt (1982a) reported 33 native species of
oval, with sparsely distributed thick bristles. Hind melyrid in New Zealand, all in subfamily DASYTINAE.
wings absent. Procoxae transverse, their cavities There are 3 described genera and 1 undescribed endemic
closed. Tibiae spinose. Tarsal segments simple. Meta- genus. Most species belong in `Dasytes' or in Arthra-
sternum short. Abdomen with 5 ventrites. canthus, the status of which is not clear.
Adults are mainly herbivorous, and may be found on
SYNOPSIS. Phycosecids are known from New Zealand, flowering shrubs, where they feed on pollen. Larvae of
New Caledonia, Vanuatu, and Australia (Lawrence & New Zealand species occur in leaf litter, in dead tree-fern
Newton 1995). The family comprises only a single genus, rachides, and under bark (Crowson 1964). Kuschel (1990)
Phycosecis, with 1 species occurring in New Zealand and collected seven species in Auckland, from vegetation,
3 others reported from Australia (Lawrence & Britton flowers, and using a Malaise trap.
1991, 1994, Watt 1982a, b, 1983a).
Adults and larvae of P. limbata are common on sandy RANGE. North ®, South ®, Stewart
beaches and coastal dunes. Both feed on small beach Kermadecs O, Three Kings ®, Chathams ®. Snares O
arthropods, and are frequently seen in dead fish or birds. Aucklands O, Campbell O, Antipodes O, Bounties O
RANGE. North ®, South ®, Stewart ®? EXAMPLE. DASYTINAE - `Dasytes' subcyaneus Broun,
Kermadecs ®, Three Kings O, Chathams ®, Snares O Ι 880 (Fig. 186).
Aucklands O, Campbell O, Antipodes O, Bounties O
REMARKS. Majer (1994) proposed a new family-group
EXAMPLE. Phycosecis limbata (Fabricius, 1781) (Fig. classification of Melyridae and related families, based on
185). cladistic analysis, in which former Dasytinae are elevated

— 45 —
to family rank. The group is badly in need of revision. dripping sap, vegetable debris, fermented fruits, on fungi,
Suggested colloquial name: flower beetles. and in stems of dead fronds of tree ferns. Many species are
known to feed on sap and fermented juices. Kuschel
SELECTED REFERENCES. Crowson (1964), Foster & (1990) recorded 16 species in 8 genera from Auckland,
Lawrence (1991), Majer (1994). from fermenting fruits and vegetables, garden litter, lawn
clippings, old logs, trunks of puriri (Vitex lucens) with
sap, dry carrion, and sooty mould on manuka (Lepto-
Superfamily CUCUJOIDEA (CLAVICORNIA) spermum scoparium). Introduced Carpophilus species
The group concept and family arrangement used here is occur in fallen fruits in orchards, and may transmit yeasts
similar to that of Pakaluk et al. (1994), Lawrence & causing fruit degradation.
Britton (1994), and Lawrence & Newton (1995). The
family name Lathridiidae is replaced by its older synonym RANGE. North ®, South ®, Stewart
Corticariidae. Members of Cucujoidea are characterised Kermadecs ®, Three Kings O, Chathams ®, Snares O
by females having a tarsal formula other than 5-5-4, Aucklands O, Campbell O, Antipodes O, Bounties O
distinctive male genital configuration, and larvae approx-
imately cylindrical, with a pygopod-like 10th abdominal EXAMPLE. NITIDULINAE - Platipidia asperella Broun,
segment (Lawrence & Britton 1994). Adults are generally 1893 (Fig. 187).
small, and most have clubbed antennae.
REMARKS. This family is very poorly known in New
Zealand, and is in need of revision.
[42] Family NITIDULIDAE Suggested colloquial name: sap beetles.
Fig. 187
SELECTED REFERENCES. Dobson (1993), Kirejtshuk
Length 2-6 mm Tarsal formula 5-5-5
(1990), Pakaluk et al. (1994).
DIAGNOSIS. Body narrowly to broadly oval, or
narrowly subparallel (Brachypeplus), moderately con-
vex to flattened, glossy to matt (Soronia), glabrous or [43] Family MONOTOMIDAE
with short and sparse pubescence, or with bristles Fig. 188, 189
(Soronia), smooth or with tubercles (Soronia). Coloration
Length 2-6 mm Tarsal formula 5-5-5
brown to black, or pale brown; elytra and often pro- or 5-5-4 (males)
notum with yellowish spots. Head usually abruptly
constricted at base of clypeus and behind eyes; fronto- DIAGNOSIS. Body narrowly oval and moderately
clypeal suture almost always absent. Antennae with an convex (Monotoma) to elongate, subparallel and sub-
abrupt, usually 3-segmented, ball-like club (Fig. 23). cylindrical (Lenax), nearly glabrous with some vestigial
Lacinia usually without a galea. Mandibles often project- pubescence to scarcely and finely pubescent. Head
ing. Pronotum transverse, often emarginated apically, abruptly constricted behind eyes, forming distinct
with anterior angles often projecting. Elytra usually temples; frontoclypeal suture absent. Antennae ap-
short, exposing pygidium and some tergites or entirely pearing 10-segmented, and antennal club appearing
covering abdomen (e.g., Platipidia), rounded apically, 2-segmented with a distinct seam between club seg-
truncate, or split with apices sinuate (some Epurea). ments (club actually might be 1-segmented with 2 distinct
Procoxae transverse, with exposed trochantins, their areas). Eyes coarsely faceted. Pronotum elongate, with
cavities open or close behind. Tibiae often expanded and anterior angles more or less produced and lateral carinae
spinose. Tarsal segments 1-3 usually lobed, and distinct. Elytra shortened, exposing at least 2 distal
segment 4 sometimes reduced. segments (Monotoma) or only the pygidium (Lenax), with
punctate striae or ridges. Procoxae globose and separate,
SYNOPSIS. Watt (1982a) recorded 20 native and 8 their cavities closed. Tarsi with the last segment en-
adventive species of nitidulid in New Zealand. We larged, as long as remaining segments combined. First
recognise 21 native and 11 adventive species (not all ventrite as long as the next 2 combined.
necessarily described) in NZAC, assigned to some 12
genera in 4 subfamilies — CARPOPHILINAE, NITIDULINAE, SYNOPSIS. New Zealand has a single native monotomid,
CILLAEINAE, and CRYPTARCHINAE. For subfamily classifi- Lenax mirandus, and 5 adventive European species of
cation see also Kirejtshuk (1986, 1990). The cosmopolitan Monotoma (Watt 1982a), all in subfamily MONOTOMINAE.
genus Carpophilus contains several species of field and The native species, originally found in Peel Forest,
stored products pests. Canterbury, has since been recorded from native forests
Adults are found in a diverse range of habitats, on both main islands. Adults of Monotoma have been
including forest growth, flowers, wounds in trees with collected mainly in the Nelson and Auckland areas.

—46—
Kuschel (1990) recorded 4 species of Monotoma from explanate, with lateral margins complete. Elytra broad
Auckland. basally, tapering posteriorly, with sides slightly ex-
Adults of Lenax mirandus have been collected from planate. Hind wings absent. Legs long. Tarsal segments
under bark of dead trees of Nothofagus fuscus and N. simple. Procoxae broadly separated (Fig. 55) and glob-
menziesii, and in dead wood and/or Platypus tunnels in ular, their cavities open.
heketara (Olearia rani). Adults of Monotoma spinicollis
in the Auckland area were found in mixed compost, lawn SYNOPSIS. Two subfamilies occur in New Zealand,
clippings, garden litter, hen-house straw, and sheep each represented by a single genus and species. Adults and
shelters in a paddock (Kuschel 1990). larvae of Agapytho foveicollis (AGAPYTHINAE) live in a
black sooty mould which grows on the honeydew excreted
RANGE. North ®, South ®, Stewart Ο by scale insects on the trunks of beech trees (Nothofagus),
Kermadecs O, Three Kings O, Chathams O, Snares O or in moss and leaf litter. Those of Priasilpha obscura
Aucklands O, Campbell O, Antipodes O, Bounties O (PRIASILPHINAE) occur in leaf litter of beech forests.

EXAMPLES. MONOTOMINAE - Lenax mirandus Sharp, RANGE. North ®, South ®, Stewart O

1877 (Fig. 188), Monotoma spinicollis Aubé, 1857 (Fig. Kermadecs O, Three Kings O, Chathams O, Snares O
189). Aucklands O, Campbell O, Antipodes O, Bounties O

REMARKS. Suggested colloquial name: none. EXAMPLES. AGAPYTHINAE —Α gapytho foveicollis Broun,
1921 (Fig. 190); PRIASILPHINAE - Priasilpha obscura
SELECTED REFERENCES. Kuschel (1979), Pakaluk et Broun, 1886 (Fig. 191).
al. (1994).
REMARKS. The status of Phloeostichidae is unsettled.
Sen Gupta & Crowson (1969b) redefined the concept of
[44] Family PHLOEOSTICHIDAE the family, discussed larval characteristics, and provided a
(= PRIASILPHIDAE)
key to the subfamilies.
Fig. 190, 191 Suggested colloquial name: bark row beetles.
Length 3-5 mm Tarsal formula 5-5-5
or (males) 5-5-4 KEY TO SUBFAMILIES OF PHLOEOSTICHIDAE
KNOWN FROM NEW ZEALAND
A diverse assemblage of primitive cucujoids, which may
be variously rearranged after adequate revision. Pakaluk et Winged, with metasternum not very short and metacoxae
al. (1994) and Lawrence & Newton (1995) listed 6 sub- narrowly separated; protrochanter partly occluded;
families of Phloeostichidae in the world fauna. The New head with a weak postocular constriction
Zealand fauna belongs to 2 subfamilies, as follows. ... (Fig. 190) .. AGAPYTHINAE
—Apterous, with metasternum very short and metacoxae
DIAGNOSIS. AGAPYTHINAE (Fig. 190): superficially sim- widely separated; protrochanter fully exposed; head
ilar to salpingids, but distinguished as follows. Body without a postocular constriction
narrow, moderately convex. Head with a distinct neck, ... (Fig. 191) .. PRIASILPHINAE
nearly as broad as maximum width of pronotum. Clypeus
large. Antennae bead-like, with a weakly defined 3- SELECTED REFERENCES. Crowson (1973), Pakaluk
segmented club. Eyes protruding. Pronotum slightly et al. (1994), Sen Gupta & Crowson (1969b).
narrower than elytra, with lateral carina incomplete
anteriorly, broadest at middle and with 2 deep foveae on
either side of base. Elytra oval, in outline, the shoulders [45] Family SILVANIDAE
slightly produced anteriorly, with 2 oblique depressions Fig. 192, 193
basally. Hind wings present. Scutellum transverse. Pro-
Length 2.5-9 mm Tarsal formula 5-5-5,
coxae separated, slightly globose and transverse. Legs appearing 4-4-4
long. Tarsal segments simple.
PRIASILPHINAE (Fig. 191): superficially similar to sil- DIAGNOSIS. Body small to medium-sized, narrowly
phids and agyrtids, but distinguished as follows. Body elongate and subparallel, slightly flattened to weakly
broadly oval and slightly depressed. Head small, par- convex, moderately glossy to matt, with inconspicuous
tially retractable into pronotum, without an apparent short, decumbent pubescence. Colour brown or rust
neck, strongly narrower than maximum pronotal brown. Head prognathous, abruptly constricted poster-
width. Antennae with a distinct 3-segmented club. Eyes iorly to form temples. Antennae usually 11-segmented,
protruding. Pronotum strongly transverse and laterally without (native species, Fig. 192) or with a 3-segmented

—47—
club (introduced species, Fig. 193); insertions often differing from that of Lawrence & Britton (1994) and
hidden from above by sides of frons. Eyes distinct, Lawrence & Newton (1995).
coarsely faceted. Pronotum usually elongate (transverse Silvanid beetles are often confused with Cucujidae/
in Ahasverus), and often with projecting front angles Laemophloeidae [46, 47], and are sometimes regarded as
and/or dentate or crenulate lateral margins; disc a subfamily of that group (e.g., Watt 1982a). Couplet I2 of
sometimes with longitudinal depressions. Elytra sub- the `quick' key should be helpful to separate them.
parallel, broader than pronotum and with distinct Suggested colloquial name: none.
shoulders, usually striately punctate. Procoxae small,
globose, narrowly separated and open or closed. Tarsal SELECTED REFERENCES. Halstead (1973, 1980),
segment 3 bilobed (exception: Brontinae), and segment 4 Lefkovitch (1961), Pakaluk et al. (1994), Thomas (1984).
usually small, giving the impression of pseudotetramerous
segmentation.
[46] Family CUCUJIDAE
SYNOPSIS. Watt (1982a) included silvanids as a subfam- Fig. 194
ily of Cucujidae [46], and reported the number of nominal Length 15-20 mm

Tarsal formula 5-5-5
species as 6 native and 6 adventive. Following Pakaluk et or 5-5-4 (males)
al. (1994) and Lawrence & Newton (1995) we recognise
silvanids and cucujids as distinct families, and find 7 DIAGNOSIS. Body elongate-oval, subparallel, strongly
native and 5 adventive silvanids represented in NZAC. In flattened, glabrous and moderately glossy. Head large,
New Zealand, silvanids are represented by 2 subfamilies, slightly broader than pronotum; labrum concealed.
BRONTINAE and SILVANINAE, each with 4 genera. Antennae 11-segmented, moniliform; basal segment
Our species of Brontinae live under bark of dead native broadly elongate, 2nd segment small and beadshaped,
trees or sometimes pine trees, or in leaf litter. Adults and 3rd one narrowly elongate and the longest, 7th and 8th
larvae of Cryptamorpha brevicornis often occur together slightly elongate, and 3 terminal segments beadshaped,
under loose bark of dead trees and in the base of dead, giving the impression of a weak club. Eyes small but
fallen leaves of nikau palm (Rhopalostylis sapida). slightly protruding. Pronotum transverse, with serrate,
Silvanids are known to feed primarily on dead plant distinct lateral carinae; front angles slightly produced
material and fungi, but our adventive species are stored anteriorly. Elytra completely covering abdomen, sub-
products pests. For instance, the cosmopolitan Ahasverus parallel in basal two-thirds, with scarcely visible striae
advena is found frequently in stored grain, nuts, and and with complete lateral carinae. Procoxae globose,
beans, where it feeds on moulds and plant debris, and broadly separated by prosternal process, with cavities
Oryzaephilus surinamensis is an economically important open; mesocoxae globose, broadly separate and open;
pest of cereals, dried fruits, and oilseeds. Adults of metacoxae transverse, separated apically but in contact
Nausibius clavicornis were introduced with Australian basally.
raisins. For details on the economically important silvanid
species, see Booth et al. (1990), Bousquet (1990), and SYNOPSIS. Cucujids are represented in New Zealand
Harney (1993). only by an undescribed species of Platisus endemic to the
Brontines may be collected by picking specimens from Three Kings Islands, first recorded by Watt (1986). Most
under bark, by sifting forest litter and processing it through specimens were collected in November and February from
Berlese funnels, or using pitfall traps in native beech or under bark of kanuka (Kunzea ericoides), especially large
kauri forests. For collection details of silvanid species old trees near mixed forest remnants.
occurring in Auckland, see Kuschel (1990).
RANGE. North O, South O, Stewart O
RANGE. North ®, South ®, Stewart O Kermadecs O, Three Kings ®, Chathams O, Snares O
Kermadecs ®, Three Kings O, Chathams O, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLE. Platisus sp. (Fig. 194).
EXAMPLES. ΒRONTINAE - Brontopriscus pleuralis REMARKS. This unique species from the Three Kings is
(Sharp, 1877) (Fig. 192); SILVANINAE - Oryzaephilus the only representative of the family Cucujidae in New
surinamensis (Linnaeus, 1758) (Fig. 193). Zealand. According to Watt (1986), representatives of
Platisus probably once occurred on the New Zealand
REMARKS. Halstead (1973, 1980) revised the genera mainland, and survived glacial climates only in the north
Silvanus and Oryzaephilus, and discussed biologically on the relatively mildly affected Three Kings, becoming
important species. Lefkovitch (1961) proposed a classi- extinct on the mainland. Several species of Platisus are
fication of genera in the two New Zealand families known from eastern Australia (Lawrence & Britton 1994).

—48—
In the past, members of the Silvanidae [45] and Laemo- EXAMPLE.
EXAMPLE. Microbrontes lineatus (Broun, 1893) (Fig.
phloeidae [47] were erroneously treated as cucujids. 195).
Suggested colloquial name: flat beetles.
REMARKS. Thomas (1995) recognised Microbrontes,
SELECTED REFERENCES. Pakaluk et al. (1994), Watt Cryptolestes, and Placonotus—males of which possess
(1986). various modifications to the antennal scape—as members
of family Laemophloeidae (Cucujidae in a broad sense).
Pakaluk et al. (1994), Lawrence & Britton (1994), and
[47] Family LAEMOPHLOEIDAE Lawrence & Newton (1995) recognised Cucujidae and
Fig. 195 Laemophloeidae as distinct families.
Suggested colloquial name: none.
Length 1.8-3 mm Tarsal formula 5-5-5
or (males) 5-5-4 SELECTED REFERENCES. Lefkovitch (1958, 1961),
Includes all the smaller, flattened cucujoids with submar- Pakaluk et al. (1994).
ginal carinae on the pronotum.

DIAGNOSIS. Body narrowly elongate, strongly flat- [48] Family PHALACRIDAE

tened, glabrous or with fine and sparse pubescence, Fig. 196
dark brown to yellowish or reddish-brown, moderately to
Length 2-3 mm Tarsal formula 5-5-5 or 5-5-4
densely punctate. Head flat, as broad as pronotum or
slightly narrower. Labrum concealed. Antennae 11- DIAGNOSIS. Body small, approximately hemispher-
segmented, moniliform, without a distinct club; males ical, flat ventrally, variably punctate, at x40 magnification
with various modifications of the scape, in Microbrontes appearing glabrous, strongly glossy, red-brown or
taking the form of a mesally directed horn or process. dark brown to black, with a metallic sheen; appendages
Eyes small. Pronotum with a sublateral carina on either paler and usually reddish-brown. Head short, broad,
side; anterior angles often acute. Elytra usually completely deeply inserted under anterior pronotal margin.
covering abdomen, striate or striate-punctate. Procoxae Mandibles moderately large. Antennae 11-segmented,
transverse, globose, their cavities open or closed; with a large 3-segmented club. Pronotum strongly
mesocoxae globose, broadly separate, open; metacoxae transverse and emarginate apically; posterior angles
transverse, broadly separate. Legs with femora swollen. tightly join-ing angular humeri of elytra in lateral view,
with lateral narrow portion of pronotum and elytra
SYNOPSIS. In New Zealand, laemophloeids are represen- explanate. Scutellum Iarge and more or less triangular.
ted by 3 adventive cosmopolitan species of Cryptolestes, 2 Elytra broadly rounded. Procoxae convex and trans-
native species (1 undescribed) in Microbrontes, and a verse, their cavities open. Tarsi 5-5-5-segmented in
third, apparently undescribed and tentatively assigned to both sexes, with the 4th segment reduced and the 3rd
Notolaemus. Microbrontes lineatus is known from several lobed (Phalacrus), or 5-5-5 in females or 5-5-4 in males,
localities in the North Island and from the Three Kings. with the 4th segment normally developed and the 3rd not
Adults of Cryptolestes pusilloides have been collected in lobed (Cyclaxyra).
Auckland, and adults of C. ferrugineus are known from
several widespread localities. SYNOPSIS. There are 2 native phalacrids in New Zea-
Most laemophloeids usually feed on moulds or on the land, with possibly a third undescribed, all in the endemic
spores and stromata of ascomycete fungi (Lawrence & subfamily CYCLAXYRINAE, and an adventive Australian
Britton 1994), and live under bark of trees, but species of subspecies in PHALACRINAE, known from Auckland and
Cryptolestes are pests of stored products – see Booth et al. Rotorua (Kuschel 1990, Thompson & Marshall 1980).
(1990), Bousquet (1990), and Harney (1993). Adults of Cyclaxyra impressa ranges from the Nelson area to the
Microbrontes lineatus have been collected from ferns at Muttonbird (Titi) Islands off Stewart Island; C. politula is
night, dead twigs of pohutukawa (Metrosideros excelsa), known from a few records on both main islands.
and nests of Turdus sp. Cryptolestes pusilloides is known Adults of Phalacrus uniformis frigoricola have been
from stored maize and wheat, and an adult specimen was found in rust fungus galls on an introduced wattle (Acacia
collected in winter from a starling (Sturnus vulgaris) nest mearnsii). Adults and larvae of Cyclaxyra impressa occur
in a bird box; C. ferrugineus has been found in stored in sooty mould growing on the honeydew of scale insects
cereals, prunes, garlic cloves, and a drying kiln. on the trunks of mountain beech and red beech (Notho-
fagus). Very little is known about C. politula, some
RANGE. North ®, South ®, Stewart O records indicating the same habitat as for C. impressa, plus
Kermadecs ®, Three Kings ®, Chathams O, Snares O moss on branches of a stunted Senecio sp. which also
Aucklands O, Campbell O, Antipodes O, Bounties O frequently bears sooty mould.

49-
-49.-
 The biology of phalacrid beetles is reviewed by Steiner and pipit (Anthus novaeseelandiae) (Watt 1980b). Adults
(1984). of Neocercus electus are known to occur in damp native
forests in the northern South Island (Watt 1980b).
RANGE. North ®, South ®, Stewart
Kermadecs O, Three Kings O, Chathams O, Snares O RANGE. North ®, South ®, Stewart O
Aucklands O, Campbell O, Antipodes O, Bounties O Kermadecs O, Three Kings ®, Chathams ®, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLE. CYCLAXYRINAE - Cyclaxyra impressa Broun,
1915 (Fig. 196). EXAMPLE. Zeonidicola chathamensis Watt, 1980 (Fig.
197).
REMARKS. The systematic position of Cyclaxyra is
controversial, and is under review by R.A. Crowson and REMARKS. It seems likely that cavognathids will be
by S.A. Slipinski (Lawrence & Newton 1995). found in nests of other bird species.
Suggested coloquial name: shining fungus beetles. Suggested colloquial name: birds' nest beetles.

SELECTED REFERENCES. Crowson (1973), Pakaluk

SELECTED REFERENCES. Pakaluk et al. (1994), et al. (1994), Sen Gupta & Crowson (1969b), Watt
Steiner (1984), Thompson & Marshall (1980). (1980b).

[49] Family CAVOGNATHIDAE [50] Family CRYPTOPHAGIDAE

Fig. 197 [assisted by R. Leschen]
Length 2.5-4.0 mm Tarsal formula 5-5-5 Fig. 198

Length 1-3.5 mm Tarsal formula 5-5-5 or (males
DIAGNOSIS. Body narrowly elongate, broadly curved of Cryptophaginae) 5-5-4
laterally, somewhat depressed, moderately glossy;
dorsal surface with decumbent pubescence; punctation DIAGNOSIS. Body oval to elongate oval, variously
irregular; colour brown to reddish-brown, often with yellow-brown or red-brown to dark brown, sometimes
paler appendages. Head narrower than pronotum; frons bicoloured, with appendages usually paler, often clothed
sometimes with impressions; frontoclypeal suture ab- with silky hairs (decumbent and/or erect), rarely glob-
sent; gular region with longitudinal grooves. Mandibles ose and appearing glabrous, convex to slightly flattened.
each with 1 or 2 cavities opening outwards. Antennae Head narrower than pronotum, without a frontoclypeal
11-segmented, with enlarged scape and with a 3-seg- suture. Antennal insertions exposed, approximated or
mented, elongate and loosely formed club. Pronotum well separated. Antennae 11-segmented, with a loose 3-
simple, approximately square or rectangular in outline, segmented club (exception: 2-segmented in Picrotus).
with margins broadly arched and angles rounded; lateral Eyes small. Pronotum subquadrate or rounded, with
carinae indistinct; disc with a smooth, raised, median lateral carinae; lateral margins occasionally serrate
impunctate area near base and with a shallow, punctate (introduced Cryptophagus); base often with a narrow,
depression on either side. Elytra with anterior angles variously formed depression; prosternal process moderately
(shoulders) slightly produced, and base concave; epi- broad, overlapping mesosternum; pronotal base usually
pleura distinct in basal third; sutural flanges widened narrower than elytral base (usually as broad as in
apically (Fig. 93), creating a small gap; striae absent. Mycetophagidae [59], or only slightly narrower). Elytra
Procoxae slightly globose and transverse, their cavities subparallel or broadly arched laterally, rounded post-
narrowly open behind. Legs stout. Protarsi and meso- eriorly; punctation confused, the punctures not in rows;
tarsi with 3 basalmost segments expanded in males. sutural flanges widened apically, forming a smalI gap;
SYNOPSIS. The cavognathids comprise 4 genera, all in epipleura incomplete, visible in basal third. Procoxae
the Southern Hemisphere. Watt (1982a) listed 2 described globose, with cavities open behind. Abdomen with 5
native species of Zeonidicola and 1 of Neocercus for New ventrites, the basal one longest.
Zealand. NZAC now has an additional 1 or perhaps 2
undescribed species of Zeonidicola. SYNOPSIS. Watt (1982a) reported 23 native and 4 ad-
Adults and larvae of Zeonidicola dumbletoni have been ventive species of cryptophagid in New Zealand. There are
found in nests of spotted shag (Stictocarbo punctatus). at present 18 native and 12 adventive species housed in
Adults of Z. chathamensis have been extracted from nest NZAC, belonging to some 12 genera in 2 subfamilies,
debris of sooty shearwater (Puffinus griseus), fairy prion CRYPTOPHAGINAE and ATOMARIINAE.
(Pachyptila turtur), giant petrel (Macronectes giganteus), Cryptophagids in general feed on hyphae, conidia, and
northern royal albatross (Diomedea epomophora sanfordi), spores of fungi, including moulds, in leaf litter, forest litter

-50-
(e.g., beech forest), vegetable debris, rotten wood, bird notha) and distinct punctate striae (irregularly punctate
and insect nests, and under bark (Leschen 1996). Cosmo- in Loberonotha). Procoxae globose, their cavities open
politan species found in stored products feed only on or closed behind. Tarsi with 4th segment reduced;
mould, their presence indicating poor storage conditions; segments 2 and 3 with setose lobes.
Bousquet (1990) reviewed stored products pests. Kuschel
(1990) recorded 13 species from Auckland, collected from SYNOPSIS. Watt (l982a) recorded 8 native languriids
heaps of garden prunings, lawn clippings, compost, hen- for New Zealand, 7 of them represented in NZAC. Our
house straw, paddocks, leaf litter, Phellinus fungus, fresh- present estimate is 7 nominal species plus 1 undescribed;
cut branches of various trees and shrubs, and foliage of these are placed in 5 genera representing 2 subfamilies —
tree ferns; some were associated with sedges and galls. XENOSCELINAE and CRYPTOPHILINAE.
Adults and larvae of Hapalips prolixus occur under
RANGE. North ®. South ®. Stewart dead leaf bases of nikau palm (Rhopalostylis sapida) and
Kermadecs O, Three Kings O, Chathams ®, Snares O in decayed tree-fern fronds. Adults of Loberus depressus
Aucklands O, Campbell O, Antipodes O, Bounties O have been collected by beating foliage of cabbage tree
(Cordyline australis), pohutukawa (Metrosideros excelsa),
EXAMPLE. CRYPTOPHAGINAE — Thortus ovalis Broun, and from tea tree blossom. Watt (1982b) recorded adults
1886 (Fig. 198). of L. nitens beaten from Coprosma repens, Carmichaelia
sp., Melicytus ramίflorus, and Pittosporum crassifolium.
REMARKS. Cryptophagids are poorly known in the Adults of Loberonotha have been collected by beating
Southern Hemisphere, and those placed in the Crypto- various vegetation, from litter, and from flowers of
phaginae (Cryptosomatulini) are in need of revision Traversia. Adults of Cathartocryptus maculosus, our sole
(Leschen & Löbl 1995). cryptophiline, have been found under bark of dead
Suggested colloquial name: silken fungus beetles. Beilschmiedia tawa and on foliage of Coprosma robusta.

SELECTED REFERENCES. Leschen (1996), Pakaluk RANGE. North ®, South ®, Stewart ®

et al. (1994). Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O

[51] Family LANGURIIDAE EXAMPLE. XENOSCELINAE — Hapalips prolixus (Sharp,

[assisted by R. Leschen] 1876) (Fig. 199).
Fig. 199
REMARKS. As at present defined, the family Languriidae
Length 2-6 mm Tarsal formula 5-5-5 includes several genera formerly placed in Cryptophagidae
This group is heterogeneous in appearance, and its classi- [50] and Erotylidae [52] (Lawrence & Newton 1995).
fication is not clear. Some members are similar to Cryp- Suggested colloquial name: slender beetles.
tophagidae [50], but are distinct in having the first ventrite
less than twice as long as the second (cf. more than twice SELECTED REFERENCES. Pakaluk et al. (1994), Sen
as long in Cryptophagidae), posterior elytral sutural gap Gupta (1968), Sen Gupta & Crowson (1969a, 1971).
absent (cf. present), epipleura complete, hind wing with 4
anal veins (cf. usually 5) and a closed radial cell, usually a
transverse groove on the anterior part of the gular region, [52] Family EROTYLIDAE
and 5-5-5-segmented tarsi in both sexes (Atomariinae of Fig. 200
Cryptophagidae have the same tarsal formula). For dis- Length 4-8 mm Tarsal formula 5-5-5
cussion see Sen Gupta (1968), Sen Gupta & Crowson
(1969a, 1971), and Lawrence & Britton (1994). DIAGNOSIS. Body elongate oval, moderately convex
DIAGNOSIS. Body moderately to strongly elongate, but with slightly flattened pronotum, usually glabrous
narrowly oval to subparallel, subcylindrical to flat- or with sparse inconspicuous pubescence, strongly
tened, pubescent or glabrous, moderately to strongly glossy, with punctation forming striae on elytra. Color-
glossy, pale brown to almost black, rarely with pale ation usually dark brown to nearly black with reddish-
maculae on elytra. Head moderately large, with eyes brown appendages, rarely yellowish brown. Head deeply
coarsely faceted. Antennae with a distinct, elongate, 3- inserted into thorax; frontoclypeal suture indistinct or
segmented club. Pronotum with lateral margins simple, absent. Antennae 11-segmented, with a 3-segmented
slightly crenulate in Hapalips and Loberonotha, usually club. Eyes prominent and coarsely faceted. Pronotum
distinctly narrower than elytral base; base sometimes with sharp lateral margins and anteriorly produced
with 2 small depressions. Elytra often elongate and front angles; base of pronotum and elytra slightly sinuate.
usually with welI defined epipleura (absent in Lobero- Elytra striate, with epipleura complete. Prosternal pro-

-51-
cess broad. Procoxae globose, inserted into articular SYNOPSIS. Hudson (1934) recorded 7 species of both-
depressions, their cavities closed. Legs robust. Tarsi riderid from New Zealand. Five are represented in NZAC,
with all segments except the last lobed and setose belonging to the subfamilies ANOMMATINAE, with 1 adven-
beneath, or simple. Femoral lines sometimes present on tive species, and BOTHRIDERINAE, with 4 native species. A
basal ventrite, which is 1.5× as long as ventrite 2. review of the world Bothriderini is provided by Slipinski
et al. (1989).
SYNOPSIS. Watt (1982a) reported 9 native species of All our bothriderids are predacious. Adults of the ad-
erotylid. Seven species are represented in NZAC, belong- ventive hypogeal species Anommatus duodecimstriatus
ing to 2 genera of DACNINAE. are recorded from rotten wood buried in earth, under
Both adults and larvae are in general mycetophagous, deeply buried stones (Sen Gupta & Crowson 1973), and
feeding within fruiting bodies of mushrooms and bracket from deep pitfall traps sealed off at the top (Kuschel 1990).
fungi. Kuschel (1990) reported two species of Crypto- Adults of Ascetoderes paynteri from the Chatham Islands
dacne from Auckland, collected from moist decayed wood have been collected in bush at night, suggesting a noct-
and from wood mould of Vitex. We have seen specimens urnal habit; others were taken from rotting Plagianthus
from dead, standing Corynocarpus laevigatus, from under logs. Adults of A. obsoletus are occasionally found under
various logs, in the dead bole of a bat roosting tree, from bark of dead tree trunks.
Piptopterus portentosus on Nothofagus, from Ganoderma
on a dead standing tree, and from Grifola colensoi. RANGE. North ®, South ®, Stewart O
Kermadecs O, Three Kings O, Chathams ®, Snares O
RANGE. North ®, South ®, Stewart ® Aucklands O, Campbell O, Antipodes O, Bounties O
Kermadecs O, Three Kings O, Chathams ®, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLES. ANOMMATINAE - Anommatus duodecim-
striatus (Müller, 1821) (Fig. 201); BOTHRIDERINAE —
EXAMPLE. Dacninae — Thallis polita White, 1846 (Fig.
Ascetoderes obsoletus (Broun, 1895) (Fig. 202).
200).

REMARKS. Suggested colloquial name: pleasing fungus REMARKS. Adult bothriderids superficially resemble
beetles. some Colydiidae [65], from which they may be distin-
guished by having exposed antennal insertions.
SELECTED REFERENCES. Pakaluk et al. (1994). Suggested colloquial name: none.

SELECTED REFERENCES. Sen Gupta & Crowson

[53] Family BOTHRIDERIDAE (1973), Slipinski et al. (1989).
Fig. 201, 202

Length 2-7 mm Tarsal formula 4 4 4 (Ascetoderes),
3-3-3 (Anommatus) [54] Family CERYLONIDAE
(= CERYLIDAE)
DIAGNOSIS. Body elongate-elliptical, subcylindrical,
sometimes with pronotum slightly flattened, glabrous, Fig. 203
slightly glossy, dark brown to black. Head narrower Length 2-3 mm Tarsal formula 4 4-4
than pronotum. Antennae 11-segmented, the basal
segment enlarged, and with a 2-segmented club; in- DIAGNOSIS. Body narrowly elongate, subparallel
sertions exposed. Eyes smalI and projecting or reduced (Philothermus) or broadly oval (Hypodacnella), moder-
(Anommatus). Pronotum trapezoidal, with front angles ately convex to flattened, glabrous to microsetose,
usually slightly produced, coarsely punctate, and often moderately to strongly glossy, often with a metallic shine
with medial impression(s); base usually distinctly nar- (Hypodacnella), pale brown or dark brown to black, with
rower than elytral base. Elytra with rounded shoulders, fine or coarse punctures. Head without a neck, partially
broadly arcuate laterally, broadest in basal half, with concealed by pronotum; frontoclypeal suture present or
longitudinal striae and coarse punctation; alternate in- absent (Philothermus). Antennae 10- or 11-segmented,
tervals costate at least apically, even intervals usually with a 1- or 2-segmented club (superficially appearing 3-
smooth; epipleura present. Procoxae globose, their segmented in Hypodacnella). Pronotum transverse or
cavities open (Anommatus) or closed (Ascetoderes); slightly elongate, distinctly margined laterally, often
metacoxae globose, moderately to widely separated. Legs coarsely punctate; base broadly arcuate or sinuate in
elongate, stout. Tarsal segments simple. Abdomen with Hypodacnella. Elytra usually striate-punctate. Hind
basal ventrite elongate, at least as long as the 2 fol- wings often with anal cell closed. Procoxae smalI and
lowing ventrites combined. semiglobular externally, their cavities open or closed;

-52-
metacoxae widely separated. Abdomen with segments DIAGNOSIS. MYCETAEINAE (Mycetaea, Fig. 204). Body
freely articulated; 1st ventrite markedly longer than small (length 1.8-2.0 mm), ovate and slightly tapering
2nd ventrite; femoral lines usually present on 1st posteriorly, moderately convex, yellow, with long and
ventrite and/or metasternum. Last ventrite with post- sparse, decumbent and erect hairs. Head partly
erior edge crenulate in Philothermus. concealed by pronotum; frontoclypeal suture distinct.
Antennae 11-segmented, with an elongate 3-segmented
SYNOPSIS. Watt (1982a) reported 5 native and 1 adven- club. Pronotum transverse, almost as broad as elytral
tive species of cerylonid from New Zealand. We now base; front angles slightly produced anteriorly; lateral
recognise 5+ native species in NZAC, belonging to 2 margins sharply carinate, with an additional lengthwise
genera in 2 subfamilies, EUXESTINAE and CERYLONINAE. ridge on either side. Elytra with punctures forming
Cerylonids occur in rotten wood, leaf litter, forest rows. Procoxae small, hemispherical, narrowly separ-
debris, and fungus-infested bark (Slipinski 1988), where ated by prosternal process, their cavities open. behind.
they probably feed on fungal hyphae and spores (Law- Abdomen with 1st sternite as long as the following 2
rence & Britton 1994). Kuschel (1990) reported Hypodac- combined, or somewhat longer.
nella rubriceps (as Hypodacne) from Auckland, collected HOLOPARAMECINAE (Holoparamecus, Fig. 205). Body
in rich, moist humus under logs, in hollow trees, and in minute (length 1.0-1.6 mm), narrowly oval, round pos-
decayed wood. We have seen specimens collected from teriorly, moderately convex, reddish brown, yellowish,
forest litter, rotten logs, and fungus on a Beilschmiedia or brown, appearing almost glabrous but with short,
log, and reared from Helichrysum coralloides. Species of inconspicuous decumbent hairs, strongly glossy. Head
Philothermus occur under bark of native trees, or in rotten somewhat concealed basally by pronotum; frontoclypeal
logs and forest litter, especially with mould. suture present. Antennae 11-segmented, with a large,
2-segmented club. Pronotum restricted behind to a
RANGE. North ®, South ®, Stewart ® distinct `waist,' with a series of laterally elongated
Kermadecs O, Three Kings O, Chathams ®, Snares O foveae near posterior margin forming a basal impression;
base markedly narrower than elytral base; additional long-
Aucklands O, Campbell O, Antipodes O, Bounties O
itudinal carinae absent. Elytra oval, with 2 impressed
lines along suture; punctate striae absent. Procoxae
EXAMPLE. EUXESTINAE — Hypodacnella rubriceps (Reit- small, hemispherical, narrowly separated by prosternal
ter, 1880) (Fig. 203). process, their cavities appearing narrowly closed be-
hind. Abdomen with 1st ventrite elongate, as long as the
REMARKS. Some cerylonids may be confused with following 3 combined.
Colydiidae [65], from which they differ, besides other
characters, in having the upper body surface glabrous or SYNOPSIS. Endomychids are represented in New Zea-
microsetose and glossy (cf. either dull or roughly sculp- land by an adventive European species of MYCETAEINAE
tured), the maxillary and labial palps geniculate (cf. never and at least 6 native species of HOLOPARAMECINAE, 3 of
so), the 1st ventrite markedly longer than the 2nd, and all them in Holoparamecus and the remainder in an undet-
5 ventrites movably articulated (cf. 1st ventrite not mark- ermined genus. Watt (1969) classified Holoparamecus in
edly longer, and 3 or 4 basalmost ventrites connate in subfamily Merophysiinae of the Corticariidae [58], and
Colydiidae). Slipinski (1988) provided characteristics of considered Mycetaea lucidus to be a synonym of Holo-
the genera of Cerylonidae. This group is in need of re- paramecus tenuis. A new classification is proposed by
vision. Pakaluk et al. (1994) and Lawrence & Newton (1995).
Suggested colloquial name: waxen beetles. Our holoparamecines occur in forest litter, organic litter
probably associated with nest debris, wood mould, and
SELECTED REFERENCES. Pakaluk et al. (1994), Sen bird nest litter; some adults have been reared from bracket
Gupta & Crowson (1973), Slipinski (1990). fungus growing on dead roots, and others from dead
Coprosma robusta. Species of this group most likely feed
mainly on moulds associated with decomposing organic
[55] Family ENDOMYCHIDAE material. Kuschel (1990) reported 2 species of Holo-
(incl. former MEROPHYSIIDAE) paramecus from Auckland, collected from logs, stumps
Fig. 204, 205 covered with fruiting bodies of larger fungi, rotted wood,
moulds inside hollow trees, humus under logs and at the
Length 1-2.0 mm Tarsal formula 4 4 4 (Mycetaea), base of herbaceous plants, and stream-bed litter.
3-3-3 (Holoparamecus)
New Zealand species of this family belong to two genera in RANGE. North ®, South ®, Stewart O
separate subfamilies, which are more readily diagnosed Kermadecs O, Three Kings ®, Chathams O, Snares O
separately, as follows. Aucklands O, Campbell O, Antipodes O, Bounties O

-53-
EXAMPLE. MYCETAEINAE - Mycetaea subterranea (Fab- ture, and 6 are known to be well established. In general,
ricius, 1801) (Fig. 204); HOLOPARAMECINAE - Holopara- members of the subfamily Coccinellinae usually feed on
mecus tenuis Reitter, 1879 (Fig. 205). aphids and homopterans, and occasionally on larvae of
Chrysomelidae and other Coccinellidae, whereas the
REMARKS. The higher classification of Endomychidae Scymninae feed on aphids, scale insects, mealybugs,
was discussed by Lawrence (1982), Slipinski (1990), and whiteflies, and plant mites (Booth et al. 1990). Kuschel
Slipinski & Pakaluk (1992). (1990) listed 26 coccinellid species from Auckland, most
Suggested colloquial name: waisted mildew beetles. of them in genus Rhyzobius, and the majority collected
from garden plants and foliage of shrubs and trees. Very
common in Auckland is the introduced Australian species
SELECTED REFERENCES. Pakaluk et al. (1994), Watt Halmus chalybeus, steel-blue in colour, and occurring in
(1969). gardens and native bush.
Collecting: sweeping and beating vegetation.

[56] Family COCCINELLIDAE RANGE. North ®, South ®, Stewart

Fig. 206 Kermadecs ®, Three Kings ®, Chathams ®, Snares O
Length 1-7 mm Tarsal formula 4-4-4 or 3-3-3 Aucklands ®, Campbell ®, Antipodes ®, Bounties O

DIAGNOSIS. Body narrowly to broadly ovate or EXAMPLE. COCCINELLINAE - Coccinella leonina Fabri-
hemispherical, moderately to strongly convex, with cius, 1792 (Fig. 206).
ventral surface usually flat, glabrous or pubescent,
slightly to strongly glossy or matt. Setation short and REMARKS. Species introduced for biological contol of
decumbent or long and protruding, often yellowish or hemipteran pests are in Coccinella, Cryptolaemus, Exo-
greyish. Coloration usually uniformly brown in smalI chomus, Halmus, Hippodamia, Illeis, Rhyzobius, Rodalia,
species and bicoloured in larger ones, variously pale and Scymnus. The family is in need of taxonomic revision.
brown, brown, yellowish, reddish brown, black, or met- Suggested colloquial name: ladybirds.
allic blue, often with spots on pronotum and elytra or with
a paler margin on pronotum and spots on elytra, the spots SELECTED REFERENCES. Bielawski (1976), Klaus-
yellow, reddish yellow, reddish brown, or black. Head nitzer (1970), LeSage (1991 c), Pakaluk eta!. (1994), Pope
partially or (rarely) almost entirely inserted into (1989), Read (1965), Sasaji (1968).
prothorax; frontoclypeal suture absent. Maxillary
palps with apical segment usually enlarged, securiform
or oblong, and apex obliquely truncated. Antennae [57] Family CORYLOPHIDAE
usu-ally 10- or 11-segmented, short, with a weak and Fig. 207
usually 3-segmented club. Pronotum transverse, usually
Length 0.8-2 mm Tarsal formula 4-4-4 or 3-3-3
strongly convex; lateral margins entire; anterior margin
often strongly emarginate; scutellum small to large. DIAGNOSIS. Body minute, oblong to broadly ovate or
Elytra usually strongly rounded, with confused punc- nearly hemispherical, moderately to strongly convex,
tation, never striate; lateral margins entire; epipleura sparsely pubescent or glabrous, often glossy. Coloration
sometimes foveate to receive femora. Procoxae trans- pale brown, brown, yellowish with brown markings, red-
verse, their cavities open. Tarsi pseudotrimerous or dish brown, or reddish yellow. Head small, completely
trimerous, with segments 1 and 2 strongly produced concealed by expanded pronotum forming a hood
beneath. Abdomen with 5 or 6 ventrites, the basalmost anteriorly (partially concealed in Orthoperus); fronto-
2 often connate; basal ventrite with impressed femoral clypeal suture absent. Antennae 9-11-segmented, with
lines (Fig. 92). a 3-segmented club. Pronotum expanded anteriorly
into a sharp margin over head, strongly transverse,
SYNOPSIS. Watt (1982a) reported 27 described native broadly rounded; posterior angles sometimes strongly
species of coccinellid and 14 adventive or introduced. produced posteriorly (Anisomeristes) or slightly conically
There are at present 22 native species and about 18 elongate (Arthrolips). Elytra subparallel or narrowed
representing introductions in NZAC, classified into some posteriorly and usually truncate, exposing pygidium. Pro-
19 genera in 4 subfamilies — COCCIDULINAE, SCYMNINAE, coxae transverse or globose, their cavities closed. Tarsi
CHILOCORINAE, and COCCINELLINAE. with 2nd segment bilobed, 3rd segment vestigial and
Most known adults and larvae are predators, feeding inconspicuous. Abdomen with 6 ventrites, the basal-
generally on soft-bodied insects and mites. A considerable most Yery long and bearing subparallel femoral lines.
number of exotic species have been introduced into New
Zealand for biological control of insect pests in agricul- SYNOPSIS. Watt (1982a) recorded 5 native and 1 adven-

-54-
tive species of corylophid in New Zealand, but the actual (exception: Melanophthalma), often with ridges and/or
number is much higher, with many species undescribed. depressions, or evenly convex and punctate, sometimes
They belong to 8 genera in 3 subfamilies — CORYLOPHINAE, with a single basal depression; lateral margins smooth or
SERICODERINAE, and PARAMULINAE. Kuschel (1990) repor- with fine dentition (e.g., Corticaria). Elytra elongate
ted 20 species in 8 genera from Lynfield, Auckland alone. oval, broadest at middle, usually much broader than
Corylophids are known to feed on fungi, and may be pronotum, or oval, sometimes swollen (Melanophthalma)),
found in association with decaying plant material such as with more or less distinct punctate striae. Procoxae pro-
forest litter, organic litter generally, compost heaps, birds' jecting, their cavities closed behind. Tarsal segments
nests, grass cuttings, mouldy hay, rotten wood, dead simple. Abdomen with 5 or 6 ventrites, the basal one
branches, litter in coastal cliff vegetation, and leaf litter at markedly elongate (much longer than the 2 following
the base of sedges and flax. Some New Zealand species ventrites combined).
have been collected from under bark, on mossy vegetation
at night, on logs overgrown with moss, on moss and liver- SYNOPSIS. Hudson (1934) recorded 57 nominal species
worts on damp slopes near streams, on foliage of native of corticariid for New Zealand. Watt (1982a) recorded 40
shrubs, occasionally on flowers, and on bracket fungi. native and 11 adventive species, and some 53 native
Sifting organic material and processing it through species and 11 adventive are now represented in NZAC.
Berlese funnels is the most efficient method of collecting; Our corticariid fauna is rich, with many undescribed
for details see Kuschel (1990). species, belonging to some 12 genera in 2 subfamilies,
LATHRIDIINAE and CORTICARΙINAE. Kuschel (1990) listed 32
RANGE. North ®, South ®, Stewart ® species in 9 genera from Auckland.
Kermadecs ®, Three Kings ®. Chathams ®, Snares O Lathridiines are subglabrous, with separated procoxae,
Aucklands O, Campbell ®, Antipodes ®, Bounties O often with carinae on the pronotum and elytra, and waxy
exudate on various parts of the body; corticariines are
EXAMPLE. Sericoderinae — Anisomeristes sharpi Mat- usually setose, with contiguous procoxae, lacking carinae
thews, 1886 (Fig. 207). on the pronotum and elytra, and with no waxy exudate.
Adults and larvae feed on fungal spores. They occur in
REMARKS. Broun (e.g., 1886, 1893a) habitually treated leaf litter, decaying trunks and stumps, wood mould, cut
branches, foliage of various plants, bracket fungi, moss
corylophids as Coccinellidae [56]. Corylophids are poorly
and liverworts on tree trunks, hollows of old trees, bush
known, and the subfamily classification is not settled,
floor litter, epiphytes, lawn clippings, and garden com-
although Lawrence & Newton (1995) have listed 4 sub-
post. Some cosmopolitan species are associated with
families in the world fauna. The group is in need of taxo-
nomic revision. stored products, and Aridius nodifer, recorded from a wide
variety of mouldy vegetables, is now established in New
Suggested colloquial name: hooded beetles.
Zealand. For other cosmopolitan species of economic
importance see Booth et al. (1990). For details on the
SELECTED REFERENCES. Endrödy-Younga (1964), bionomics of species occurring in Auckland, see Kuschel
Gressitt & Samuelson (1964b), Pakaluk et al. (1994). (1990).

RANGE. North ®, South ®, Stewart ®

[58] Family CORTICARIIDAE Kermadecs ®, Three Kings ®, Chathams ®, Snares O
(= LATHRIDIIDAE = LATRIDIIDAE) Aucklands ®, Campbell ®, Antipodes O, Bounties O
Fig. 208
Length 0.8-2.2 mm Tarsal formula 3-3-3 EXAMPLE. LATHRIDIINAE - Enicmus caviceps Broun,
(1893) (Fig. 208).
DIAGNOSIS. Minute oblong beetles, with pronotum
usually markedly narrower than elytra (as broad as REMARKS. The name Corticariidae has priority over the
elytral base, or nearly so in Metophthalmus (= Lithostyg- universally used Lathridiidae for this family (Pakaluk et
nus)). Body surface glabrous or finely tomentose, often al. 1994). First revisions of New Zealand corticariid spec-
covered by a waxy exudate, moderately glossy to matt, ies were published by Belon (1884, 1897). Watt (1969)
smooth and punctate or with ridges, tubercles, and/or provided a generic key to New Zealand Corticariidae,
depressions. Head with clypeus laterally expanded including Holoparamecus, now referred to Endomychidae
in front of antennal insertions; frontoclypeal suture pres- [55].
ent. Antennae 11-segmented, with a 2- or 3-segmented Suggested colloquial name: mildew beetles.
club; scape usually enlarged, often globular. Eyes small
but protuberant, sometimes reduced, coarsely faceted. SELECTED REFERENCES. Belon (1884, 1897), Gressitt
Pronotum usually distinctly narrower than elytra at base & Samuelson (1964a), Pakaluk et al. (1994), Watt (1969).

-55-
Superfamily TENEBRIONOIDEA (HETEROMERA) [60] Family ARCHEOCRYPTICIDAE
Members of this group are distinct in having tarsi 5-5-4- Fig. 210
segmented in both sexes, rarely 4-4-4 in a few lineages,
Length 2.7-3.5 mm Tarsal formula 5-5-4
trochanterofemoral attachment strongly oblique (Fig. 59),
procoxae tending to be conical and projecting, hind wings DIAGNOSIS. Body uniformly oval, slightly flattened,
with a reduced number of veins (4 or fewer) behind the dark brown to almost black, with short pubescence. Head
media posterior (MP), aedeagus heteromeroid with the partially retracted into prothorax, broadly arcuate
tegmen incomplete ventrally, and in certain characteristics anteriorly; frontoclypeal suture distinct. Maxillary
of the larvae (Lawrence & Britton 1991, 1994). palps with apical segment enlarged and truncate.
Antennae 11-segmented, with a 3-segmented club. Eyes
coarsely faceted. Pronotum transverse, with sharp
[59] Family MYCETOPHAGIDAE lateral margins, slightly emarginate anteriorly and in-
Fig. 209 significantly sinuate basally. Elytra arcuate laterally, with
Length 1.5-3 mm

Tarsal formula 4-4-4 (females), confused punctation. Procoxae globose, separated by
3-4-4 (males) prosternal process, which is gradually expanded api-
cally. Tarsal segments simple. Abdomen with 5 vent-
DIAGNOSIS. Body broadly oval, somewhat flattened, rites, the first 2 connate (3 connate in Tenebrionidae
evenly pubescent, moderately glossy, punctate, yellowish [68]); basal ventrite slightly longer than 2nd ventrite.
to dark brown to black, sometimes with pale or dark
markings on elytra and/or pronotum. Head narrower SYNOPSIS. There is a single adventive species of arch-
than pronotum; antennal insertions exposed. Maxillary eocrypticid, Archeocrypticus topali, in New Zealand.
palps with last segment enlarged. Antennae 11- Adults and larvae are reported from leaf litter and some
segmented, usually with a 3-segmented club. Eyes polypore fungi with softer fruiting bodies (Lawrence &
coarsely faceted. Pronotum transverse; sides arcuate, Britton 1994). Our species has been recorded from gar-
sometimes with minute serrations, smoothly continuous dens, parks, litter in beech forest, under stones (rarely),
with sides of elytra; base as wide as elytra. Scutellum under seaweed on beaches, moss on banks, nests of gulls
transverse. Elytra broadly oval, punctate. Procoxae (Larus sp.), on mummified apples, and under mature
transverse, their cavities open. Legs well developed. cabbages. In all these habitats it is probably associated
with moulds or other fungi. Many adults and larvae have
SYNOPSIS. Watt (1982a) reported 12 native and 1 ad- been collected in pitfall traps in pastures around Nelson.
ventive species of mycetophagid for New Zealand. These
are in 3 genera of subfamily MYCETOPHAGINAE. Kuschel RANGE. North ®, South ®, Stewart O
(1990) recorded 5 native species and 3 adventive from Kermadecs O, Three Kings O, Chathams O, Snares O
Lynfield, Auckland. Aucklands O, Campbell O, Antipodes O, Bounties O
Adults and larvae may be found on or in fungi and
organic material infested with fungi. Some species have EXAMPLE. Archeocrypticus topali Kaszab, 1964 (Fig.
been collected by beating or sweeping foliage of native 210).
shrubs. The adventive species have been found in a sheep
carcass, coarse prunings, a starling nest, bumble bee nests, REMARKS. Suggested colloquial name: ancient fungus
lawn clippings, compost, trees, shrubs, and a rotting seed beetles.
head of Cynara scolymus (Kuschel 1990).
SELECTED REFERENCES. Kaszab (1981, 1984), Law-
RANGE. North ®. South ®, Stewart O rence (1994), Watt (1974a).
Kermadecs O, Three Kings ®, Chathams ®, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
[61] Family CIIDAE
EXAMPLE. MYCETOPHAGINAE — Triphyllus hispidellus Fig. 211
(Broun, 1880) (Fig. 209). Length 1-3 mm

Tarsal formula 4 4 4
REMARKS. Mycetophagids are poorly known in New DIAGNOSIS. Body elongate, subcylindrical or narrow-
Zealand. The European concept of Triphyllus applied to ly ovate and moderately convex, yellowish, brownish,
our native species needs clarification. reddish brown, dark brown, or greenish, uniform in colour
Suggested colloquial name: hairy fungus beetles. or with yellowish or brownish elytral spots sometimes of
irregular shape, glabrous and strongly glossy or mod-
SELECTED REFERENCES. Lawrence (1987), Leschen erately glossy and with short decumbent hairs and/or
& Lawrence (1991), Parsons (1975). bristles. Head deflexed, with a distinct frontoclypeal

—56—
ridge, this in male sometimes forming a plate or 2 elongate at least in mesotarsus and metatarsus, and
tubercles or horns. Maxillae with reduced lobes and penultimate segment lobed. Abdominal ventrites mov-
fusiform palps. Antennae 8-10-segmented, with usually able (not fused).
a 3-segmented loose club (Fig. 9) (cf. a similar 3-seg-
mented compact club in Curculionidae: Scolytinae [82]). SYNOPSIS. Watt (1982a) reported 38 native species of
Pronotum large, partially or entirely covering head melandryid for New Zealand. The species are assigned to
(Fig. 9); base often approximately straight; lateral edges 9 genera, 5 in subfamily MELANDRYINAE and the remainder
incomplete. Elytra elongate, usually subparallel, with of uncertain affiliation. Kuschel (1990) recorded 8 species
punctures confused or in rows. Procoxae transverse, in 7 genera from Auckland.
projecting, their cavities open. Tibiae often with spines Adult melandryids occur in rotten wood, logs, organic
on outer edge. Abdomen with 5 ventrites; basal ventrite litter including leaf litter, cut branches, flowers, at the
usually with a setose fovea in male. base of bush-floor sedges, and on forest foliage. Adults
are very agile, and are capable of jumping; some have been
SYNOPSIS. Watt (1982a) recorded 20 native species of collected using interception traps. Larvae inhabit hard
ciid for New Zealand. These belong to 5 genera of substrates, such as bracket fungi or wood (Lawrence &
subfamily CIINAE. Kuschel (1990) recorded 17 species Newton 1995).
from Lynfield, Auckland, in 5 genera.
Adults and larvae are typically mycetophagous in fruit- RANGE. North ®, South ®, Stewart
ing bodies of shelf fungi (Polyporaceae). Some occasion- Kermadecs O, Three Kings ®, Chathams ®, Snares
ally occur in forest litter, dead wood, or cut branches. Aucklands ®, Campbell ®, Antipodes ®, Bounties O

RANGE. North ®, South ®, Stewart ® EXAMPLE. MELANDRYINAE- Hylobia nubeculosa Broun,

Kermadecs O, Three Kings ®, Chathams ®, Snares O 1886 (Fig. 212).
Aucklands O, Campbell O, Antipodes O, Bounties O
REMARKS. Suggested colloquial name: leaping beetles.
EXAMPLE. CIINAE - Cis zeelandicus Reitter, 1880 (Fig.
211). SELECTED REFERENCES. Crowson (1966b), Gressitt
& Samuelson (1964a), Viedma (1966).
REMARKS. Suggested colloquial name: shelf fungus
beetles.
[63] Family MORDELLIDAE
SELECTED REFERENCES. Abdullah (1973), Lawrence Fig. 213
(1971), Zimmerman (1938).
Length 4-17 mm Tarsal formula 5-5-4
DIAGNOSIS. Body wedged-shaped, tapered posteri-
[62] Family MELANDRYIDAE orly, laterally compressed, characteristically humped
Fig. 212 in lateral view, with the last abdominal segment
Length 1.5-12 mm Tarsal formula 5-5-4 produced and forming an acute process extending
beyond elytra (often extremely elongate); surface with
DIAGNOSIS. Body elongate, approximately wedge- fine, decumbent pubescence; coloration brown with
shaped, tapering posteriorly, convex dorsally and darker markings on pronotum, uniformly black, or black
ventrally, with fine and decumbent pubescence, finely with small white spots on elytra and often on lateral parts
or coarsely punctate; coloration uniformly dark brown to of sternites. Head deflexed, flat ventrally and concealing
black, pale brown, or yellowish orange, sometimes with prosternum (in resting position), abruptly constricted
yellow, orange, or brown markings on elytra and pro- behind eyes and forming a narrow neck. Maxillary
notum. Head deflexed, deeply inserted into prothorax, palps with last segment expanded. Antennae 11-
not constricted behind. Maxillary palps with last seg- segmented, filiform or slightly serrate. Eyes entire.
ment enlarged, often securiform. Antennae usually Pronotum as broad as elytra or slightly broader,
filiform or incrassate, but without a distinct club. Eyes transverse, sinuate basally and with complete lateral
entire, not pubescent. Pronotum with lateral carinae carinae. Elytra tapering posteriorly. Procoxae projec-
incomplete anteriorly, sometimes with small longitudinal ting, their cavities open; metacoxae enlarged, plate-
depressions. Elytra tapering posteriorly, sometimes like. Tibiae often with comb-shaped spines.
with impressed longitudinal lines subparallel to suture.
Procoxae projecting, their cavities open behind. Legs SYNOPSIS. Watt (1982a) recorded 5 native species of
long. Tibial spurs often comb-shaped, extremely elon- mordellid for New Zealand. Kuschel (1990) listed 6
gate on metatibiae (Fig. 84). Tarsi with basal segment species, including 1 adventive, in 4 genera from Lynfield,

—57—
Auckland. Two genera are in subfamily MORDELLINAE and cocoon of a cerambycid beetle. Kuschel (1990) reported
the remainder are unassigned. Allocinops brookesi to be a predator on larvae of Oemona
Adults have often been found on flowers of manuka hirta (Cerambycidae). The larvae are known to be
(Leptospermum scoparium), ragwort (Senecio jacobaea), entomophagous parasitoids of immature Hymenoptera,
umbelliferous flowers, foliage of native shrubs and trees, Coleoptera (Cerambycidae), and Blattodea, and their
forest litter, and grass. Larvae occur in rotten wood or development involves both external and internal feeding
stems of various species, e.g., castor oil plant (Ricinus stages (Selander 1991). Unlike other rhipiphorids, the
communis). larva of Rhipistena is an eruciform, apparently free-living
Collecting: sweeping or beating foliage of trees, shrubs, predator of larvae of Prionoplus (Cerambycidae) (Hudson
and lower vegetation, or from interception traps. 1934).

RANGE. North ®, South ®, Stewart RANGE. North ®. South ®, Stewart O

Kermadecs O, Three Kings O, Chathams ®, Snares O Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O Aucklands O, Campbell O, Antipodes O, Bounties O

EXAMPLE. MORDELLINAE — Mordella antarctica White, EXAMPLE. PELECATOMINAE— Rhipistena lugubris Sharp,
1846 (Fig. 213). 1878 (Fig. 214).

REMARKS. New Zealand's mordellids are currently REMARKS. The relationships of our rhipiphorid genera
being studied by Dr M.E. Franciscolo of Genoa, Italy. in the world context require investigation.
Suggested colloquial name: pintail beetles. Suggested colloquial name: antlered beetles.

SELECTED REFERENCES. Batten (1990), Ermisch SELECTED REFERENCES. Selander (1991).

(1950), Franciscolo (1980), Odnosum (1991).
[65] Family COLYDIIDAE
[64] Family RHIPIPHORIDAE Fig. 215, 216
Fig. 214 Length 1-19 mm

Tarsal formula 4-4-4
Length 6-8 mm Tarsal formula 5-5-4
DIAGNOSIS. Body broadly oval or ovate to narrowly
DIAGNOSIS. Body elongate, more or less laterally subparallel, sometimes extremely elongate (e.g.,
compressed, tapering posteriorly but lacking an Rc
ytinous), onvex to flattened, often with ridges, protuber-
abdominal spine, finely pubescent, the pubescence dark ances, or ornamentation, dark in colour, usually brown
or yellowish and decumbent; coloration pale brown to to black, sometimes variegated with paler or darker mark-
dark brown, with head sometimes black and elytra usually ings, usually matt with various pubescence, decumbent
paler. Head deflexed (in natural position), abruptly or erect or with bristles or scales, rarely glossy and
constricted behind eyes to form a neck. Antennae with subglabrous. Head usually partially retracted into tho-
last 7 segments flabellate in males (Fig. 33). Eyes rax, often with a well developed frontoclypeal ridge
usually emarginate, more or less kidney-shaped and which may be expanded forwards; antennal insertions
partly surrounding base of antennae. Pronotum trapez- usually concealed. Maxillary palp with last segment
oidal, sinuate basally. Elytra covering abdomen entirely, small. Antennae 10- or 11-segmented, incrassate, with
or shortened and exposing at least abdominal apex, with a 2- or 3-segmented club. Pronotum usually transverse,
longitudinal ridges but not striate. Procoxae projecting, occasionally quadrate or elongate, usually explanate
their cavities open behind. Legs slender. Tibial spurs laterally, with lateral carinae complete and often
non-serrate. Tarsal segments simple; tarsal claws crenulate or dentate, sometimes expanded laterally
toothed (Fig. 87). and with apically produced front angles; disc usually
grooved, carinate, and/or tuberculate. Elytra with ridges
SYNOPSIS. Hudson (1934) listed 5 native species of and/or tubercles and rows of punctation. Procoxae
rhipiphorid for New Zealand, but Watt (1982a) recorded globular, their cavities open or closed. Tarsal segments
only 4. The species are placed in 3 genera of subfamily simple. Abdomen with 5 ventrites, the basal 3 free or
PELECATOMINAE, but the status of 1 genus is uncertain, and connate.
it may have to be synonymised. Kuschel (1990) recorded
1 species from Lynfield, Auckland. SYNOPSIS. Watt (1982a) reported 196 native species of
Adults are found on scrubby vegetation, in dead wood, colydiid for New Zealand. There are 133 species repre-
and on foliage of native shrubs, with one record from the sented in NZAC, belonging to some 26 genera in 2 sub-

— 58 —
families, PYCNOMERINAE and COLYDIINAE. The status of tuberculate, with complete epipleura. Procoxal cavities
some genera needs revision. Kuschel (1990) recorded 24 closed behind by lateral extensions of prosternal
species in 14 genera from Lynfield, Auckland. intercoxal process (Fig. 39) (in Tenebrionidae [68]
Pycnomerines are distinct by having partly closed closed by mesal extensions of pronotal hypomera — Fig.
procoxal cavities, a laterally expanded prosternal process, 38). Tarsal segments simple. Abdomen with 3 basal
and widely separated metacoxae, and by features of their ventrites connate.
larvae; colydiines are distinct by having procoxal cavities
either open or closed by projections of the notum, SYNOPSIS. Watt (1982a) reported 20 native species of
moderately widely separated metacoxae, and by larval ulodid (as Zopheridae) for New Zealand. Fourteen species
characteristics (Lawrence & Britton 1991). are represented in NZAC, belonging to 4 genera, of which
Adults may be found under bark, on standing dead tree one is confined to New Zealand.
trunks, in rotten wood, cut branches, leaf litter, forest litter, Adults and larvae of Brouniphylax squamiger occur in
and on various trees and shrubs. Most species are believed dead bracket fungi (Fomes), and adults alone are some-
to be mycetophagous, but Lawrence & Britton (1994) times found under logs. Both stages of Syrphetodes are
reported certain New Guinean species as being predators found under loose bark of standing dead trees or logs and
of platypodine weevils. Larvae are generally taken from in rotten branches; adults also are found on the underside
rotten wood. of logs. Kuschel (1990) reported B. squamiger in dead
Polyporaceae on decaying logs from Lynfield, Auckland.
RANGE. North ®. South ®. Stewart
Kermadecs ®, Three Kings ®, Chathams ®, Snares O RANGE. North ®, South ®, Stewart O
Aucklands ®. Campbell ®. Antipodes O, Bounties O Kermadecs O, Three Kings ®, Chathams O, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLES. COLYDIINAE — Pristoderus antarcticus
(White, 1846) (Fig. 215), Rhizonium antiquum Sharp, EXAMPLE. Syrphetodes marginatus Pascoe, 1875 (Fig.
1876 (Fig. 216). 217).

REMARKS. The placement of Rhizonium antiquum in REMARKS. Ulodids are superficially similar to tenebri-
Colydiidae is uncertain, and should be treated as a tem- onids [68], from which they can be readily separated by
porary arrangement. Lawrence (1994) has shown that having exposed antennal insertions, an apically expanded
Pycnomerinae are more closely related to Monommatidae prosternal process, and in larval features (clypeus fused to
and Zopheridae (sensu Lawrence & Newton 1995) than frons, and frontoclypeal suture obliterated). In tenebrionid
they are to Colydiinae. larvae the clypeus is always distinct from the frons. For
New Zealand's Colydiidae are under investigation by comparisons of Ulodidae, Chalcodryidae [67], and Zoph-
Dr M.A. Ivie (Montana State University, Bozeman, eridae see Watt (1974a), Doyen & Lawrence (1979),
U.S.A.) and by Dr S.A. Slipinski (Polish Academy of Lawrence (1994), and Lawrence & Newton (1995). For a
Science, Warsaw, Poland). key to all genera of Ulodidae see Lawrence (1994).
Suggested colloquial name: rough mould beetles. Suggested colloquial name: false darkling beetles.

SELECTED REFERENCES. Sharp (1876c), Slipinski & SELECTED REFERENCES. Doyen & Lawrence (1979),
Burakowski (1988), Wollaston (1873). Lawrence (1994), Watt (1974a).

[66] Family ULODIDAE [67] Family CHALCODRYIDAE

(formerly as ZOPHERIDAE) Fig. 218
Fig. 217 Length 5-22 mm Tarsal formula 5-5-4
Length 4-20 mm Tarsal formula 5-5-4 DIAGNOSIS. Body fairly soft, loosely articulated, nar-
DIAGNOSIS. Body broadly oval, slightly depressed, rowly elongate, subparallel, moderately convex, glossy,
brown, matt (exception: Arthopus, glabrous and strongly pale to dark green or brownish to dark brown, often
glossy); dorsal surface with decumbent and/or erect with metallic reflections, and sometimes with patches of
pubescence, bristles, or scales usually forming a pat- yellowish pubescence; surface punctate. Head prognath-
tern. Head partially retracted into prothorax; antennal ous, slightly narrowed behind eyes, with a broad neck;
insertions exposed. Antennae 11-segmented, filiform, antennal insertions exposed. Maxillary palp with last
with at most a weak club. Eyes almost circular. Pro- segment enlarged, securiform. Antennae 11-segmented,
notum narrower at base than elytra, often with front filiform, sometimes with last 3 segments slightly larger
angles produced anteriorly. Elytra usually broad, often but not forming a distinct club. Pronotum transverse,

— 59 —
rectangular, with sides explanate. Elytra subparallel, [68] Family TENEBRIONIDAE
without clearly defined striae but with irregular rows of Fig. 219-221
punctures; epipleura with a carina extending almost to
apex. Procoxae projecting, their cavities large, trans- Length 2.9-18 mm Tarsal formula 5-5-4 or
verse, closed behind partly by inward extensions of (Archaeoglenes) 4-4-4
propleura and partly by lateral extensions of narrow Adult tenebrionids are highly variable in form, and some
intercoxal process; mesocoxal cavities narrowly separated, are superficially similar to members of other families,
open laterally. Legs long and slender. Tarsal segments e.g., Carabidae [2], Scarabaeidae [15], Ulodidae [66], and
elongate, not lobed. Abdomen with 3 or 4 basal ventrites Chrysomelidae [77]. However, they are distinct in the
weakly connate. following combination of characters.

SYNOPSIS. Chalcodryids are the only endemic family in DIAGNOSIS. Body usually black or brown, rarely pale,
New Zealand, with 5 native species in 3 genera. A key to moderately glossy (rarely glossy metallic) to matt, often
genera and species is provided by Watt (1974b), together subglabrous. Head not sharply constricted behind
with a detailed description of this family. Distribution eyes; antennal insertions under a canthus (Fig. 11),
records suggest a northern limit to the south of Auckland. which conceals at least base of scape from above and
Watt (1974b) collected adults by beating branches usually extends back to anterior margin of eye, on
covered with moss and/or lichen in cool, wet forests, which it usually encroaches. Maxillary palp with last
especially of Nothofagus. Adults of Chalcodrya are en- segment often securiform. Antennae usually filiform,
countered most frequently, and their larvae have been incrassate, distinctly clubbed or bluntly serrate. Eyes
found in short refuge galleries in dead suppressed twigs or usually emarginate. Pronotum usually carinate or
small branches on various trees; they can be collected at explanate laterally. Elytra usually narrowly oval or
night from moss-covered branches (Watt 1974b). Larval subparallel, with distinct epipleura, if striate then
gut contents predominantly consist of lichen and moss, usually with 9 or fewer striae (exception: 10 striae in some
with some fragments of mites and spiders (Watt 1974b). Lagriinae), often with a scutellary striole or with sutural
striae diverging basally. Hind wings sometimes reduced.
RANGE. North ®, South ®, Stewart Procoxae variable in shape, often projecting, their
Kermadecs O, Three Kings O, Chathams O, Snares O cavities almost always closed behind by extensions of
Aucklands O, Campbell O, Antipodes O, Bounties O pronotal hypomera, but not by lateral extensions of
prosternal intercoxal process as in Ulodidae [66] and
EXAMPLE. Chalcodrya variegata Redtenbacher, 1868 Chalcodryidae [67], without exposed trochantins. Troch-
(Fig. 218). anters heteromeroid (e.g., Fig. 59). Legs variable in form
and function, from stout and specialised for digging to
REMARKS. Chalcodryids in many features are similar to slender for running. Tarsal segments usually not lobed
Periomylopidae, Ulodidae, and Tenebrionidae. They may below, the claws simple or pectinate (Alleculinae) but
be distinguished from Tenebrionidae [68] in having the never appendiculate. Abdomen with 5 ventrites, the first
procoxal cavities partly closed by lateral extensions of the 3 connate, and often with intersegmental membrane
prosternal intercoxal process, 4 connate abdominal stern- clearly visible between ventrites 3/4 and 4/5.
ites (3 in Tenebrionidae), and exposed antennal insertions.
Chalcodryids differ from Ulodidae [66] in their fairly soft SYNOPSIS. Tenebrionids are a major group, comprising
and loosely articulated body (cf. compact and strongly approximately 15 000 described species worldwide. There
sclerotised), narrow prosternal intercoxal process, and are 149 valid species in New Zealand, 10 of them adven-
lack of concealed lateral procoxal extensions. According tive, in 36 genera, 16 tribes, and 8 subfamilies (Watt 1992)
to Watt (1974b), chalcodryids differ from Periomylopidae - LAGRIINAE, ZOLODININAE, PIMELIINAE, TENEBRIONINAE,
in having concealed trochantins on the procoxae and ALLECULINAE, DIAPERINAE, and COELOMETOPINAE. Tenebri-
having usually 4 connate abdominal sternites. onids constitute 3.3% of the total known species of
Lawrence (1994) transferred the Tasmanian genus Coleoptera in New Zealand (Watt 1992). Kuschel (1990)
Sirrhas—placed in Chalcodryidae by Watt (1974b)—to recorded 21 species in 15 genera from Lynfield, Auckland.
the Periomylopidae, leaving Chalcodryidae restricted in The biology of New Zealand tenebrionids is recorded
distribution to New Zealand. Lawrence (1994) discussed by Watt (1992). Many of our native species live in rotten
the new concept of the family. wood (Aphtora, Menimus, Uloma, Ulomotypus, Zolodi-
Suggested colloquial name: southern beech beetles (R. nus), or in powdery, dry rotten wood (Mimopeus opac-
Palma, in litt.) ulus). Archeoglenes occurs in leaf litter and under logs.
Menimus inhabits dead fruiting bodies of large woody
SELECTED REFERENCES. Lawrence (1994), Watt fungi. Actizeta and Chaerodes live in sandy beaches,
(1974b). recalling the many species overseas known mainly from

—60—
arid and semi-arid habitats. Some species (in Artystona, gate, slightly depressed medially; lateral carinae in-
Pseudhelops, Partystona, and Cerodolus) feed on lichens distinct. Elytra with angular shoulders and with striae
at night. Lorelus lives in dead tissue of tree ferns and dead weakly defined except for lateral ones. Procoxae globose,
flower stalks of speargrass. Amarygmus, Chrysopeplus, their cavities closed behind; procoxal process broad;
and Demtrius occur under loose bark of standing dead both procoxae and mesocoxae widely separated. Tarsal
trees. Gonocephalum, Mimopeus, Omedes, Zomedes, and segments simple. Abdomen with 5 ventrites, the basal 2
Adeliini are frequently found under stones. Tanychilus connate.
and Xylochus occur on flowers. Mimopeus occasionally
feeds on live plant tissue. SYNOPSIS. Only a single species of prostomid, Dryo-
The adventive cosmopolitan elements in our fauna are cora howitti, is recorded from New Zealand.
in the genera Alphitobius, Gnathocerus, Tenebrio, and Adults and larvae occur in logs of rimu (Dacrydium
Tribolium, of which a few species infest stored products. cupressinum), totara (Podocarpus totara), and hinau
Collecting: hand picking from under stones or logs (at (Elaeocarpus dentatus) which have reached the red stage
night with a headlamp); processing organic litter through of decay, breaking up into rectangular pieces with mud-
Berlese funnels; pitfall trapping; and sifting sand. like substrate in between, on which they feed.

RANGE. North ®, South ®, Stewart ® RANGE. North ®, South ®, Stewart O

Kermadecs ®, Three Kings ®, Chathams ®, Snares ® Kermadecs O, Three Kings O, Chathams O, Snares O
Aucklands ®, Campbell ®, Antipodes ®, Bounties ® Aucklands O, Campbell O, Antipodes O, Bounties O

EXAMPLES. LAGRIINAE — Chaerodes trachyscelides EXAMPLE. Dryocora howitti Pascoe, 1868 (Fig. 222).
White, 1846 (Fig. 219); PIMELIINAE — Actizeta albata
Pascoe, 1875 (Fig. 220); TENEBRIONINAE — Mimopeus REMARKS. In earlier classifications prostomids were
elongatus (Brême, 1842) (Fig. 221). sometimes treated as a subfamily of Cucujidae [46], prob-
ably owing to their strongly depressed body. The genus
REMARKS. Keys to subfamilies of Tenebrionidae of the Dryocora is also known from Australia.
world (based on adults and larvae) are provided by Watt Suggested colloquial name: red log beetles.
(1974a). Keys to New Zealand subfamilies, genera, and
species of Tenebrionidae (based on adults) are provided by SELECTED REFERENCES. Pascoe (1868), Schawaller
Watt (1992). Doyen et al. (1990) discussed the subfamily (1993).
classification of Tenebrionidae, with Zolodininae treated
as a subfamily (in agreement with Watt 1974a). For cos-
mopolitan pest species see Booth et al. (1990). [70] Family OEDEMERIDAE
Adults and larvae of Tenebrionidae may be reared on a Fig. 223
diet consisting of wholemeal flour and dried yeast, with Length 6-20 mm Tarsal formula 5-5-4
access to water.
Suggested colloquial name: darkling beetles. DIAGNOSIS. Body soft, elongate, subparallel, finely
and uniformly pubescent, testaceous or fuscous, some-
REFERENCES. Doyen et al. (1990), Skopin (1964), times with dark brown pattern, or uniformly black. Head
Watt (1967, 1968, 1974a, 1988, 1989b, 1992), Watt & small, moderately deflexed, elongate before eyes, equal
Triplehorn (1991). in width to pronotum. Maxillary palp showing sexual
dimorphism in almost alI species, the apical segment
expanded and truncate in males. Antennae long, usu-
[69] Family PROSTOMIDAE ally filiform, 11-segmented. Eyes variable in size, entire
Fig. 222 or slightly emarginate near antennal insertions. Pro-
thorax subcylindrical, usually longer than broad, widest
Length 6-8 mm Tarsal formula 4 4 4 anteriorly or at middle, without lateral carinae, distinct-
DIAGNOSIS. Body narrowly elongate, subparallel, ly narrower than elytra. Elytra either elongate, with
strongly depressed, glabrous, glossy, reddish brown, fully developed hind wings, or shortened and exposing
yellowish brown, or brown, with surface punctate. Head abdomen, with reduced hind wings, rounded apically,
prognathous, large, slightly broader than pronotum at with indistinct epipleura restricted to basal half; elytral
base, strongly produced posterolaterally behind eyes, surface sometimes costate. Procoxae projecting, their
with a distinct narrow neck; frontoclypeal suture cavities open. Legs slender and long, with hind femora
present. Antennae 11-segmented, with a 3-segmented usually swollen. Tarsi with penultimate segment
elongate club; scape enlarged, with segments bead- bilobed; claws simple or toothed. Abdomen with 5
shaped. Eyes small, round, projecting. Pronotum elon- ventrites, and with genitalia usually exposed in male.

-61 —
SYNOPSIS. There are 21 species of oedemerid in New elytral width, subquadrate or subcylindrical, with lateral
Zealand, of which 18 are native and 3 adventive (Hudson margin smooth. Elytra flat to slightly convex, with larger
1975, Watt 1982a). They belong in 6 genera representing punctures bearing shorter decumbent setae and smaller
2 subfamilies, NACERDINAE and OEDEMERINAE. All our punctures bearing longer, secondary elytral setae; epi-
native species are oedemerines, and some are restricted to pleura distinct, narrow, extending almost to apex of
the alpine zone of the South Island. Of the adventive elytra. Legs long, slender. Procoxae conical, projecting,
species, 2 are Australian and the other, our only nacerdine, their cavities open posteriorly and internally; meso-
is cosmopolitan. Kuschel (1990) recorded 2 adventive coxae narrowly separated. Tarsi with penultimate and
species collected at light from Lynfield, Auckland. antepenultimate segments lobed beneath and with dense,
Adult oedemerids occur in coastal habitats, under fine ventral setae. Abdomen with 5 movable ventrites.
driftwood just above high tide, and in inland habitats on
vegetation; they may be common on flowers or herbage or SYNOPSIS. According to Watt (1987) and Pollock
under driftwood. Some species are restricted in habitat, (1995) there are 7 native species of pyrochroid described
whereas others range from coastal to inland areas (Hudson and 3 or so undescribed. They belong to 3 endemic New
1975). Larvae live in decaying and/or wet wood, some- Zealand genera of subfamily PILIPALPINAE.
times in timber partly or intermittently submerged in saline Adults are found under bark of dead trunks and
or fresh water, and are common in driftwood along coastal branches, or on foliage or flowers. Watt (1987) found very
shores; others occur in decaying Olearia and Dracophyllum little pollen in gut contents, which usually contained
logs and dead branches inland (Hudson 1975). fungal hyphae and spores and much amorphous vegetable
matter. Larvae are almost invariably subcortical on logs,
RANGE. North O, South ®. Stewart dead branches, or standing dead trunks, where they feed on
Kermadecs ®, Three Kings O, Chathams ®, Snares O decaying cambial matter and fungal hyphae. Larvae of
Aucklands O, Campbell O, Antipodes O, Bounties O various sizes are found at most times of the year, indicating
that the life cycle may take 2 years or more to complete
EXAMPLE. OEDEMERINAE - Thelyphassa lineata (Fab- (Watt 1987).
ricius, 1792) (Fig. 223).
RANGE. North ®. South ®, Stewart O
REMARKS. Haemolymph of oedemerids contains the Kermadecs O, Three Kings O, Chathams O, Snares O
irritant cantharidin, which may cause blistering of the skin, Aucklands O, Campbell O, Antipodes O, Bounties O
so caution is necessary in handling them.
Suggested colloquial name: lax beetles. EXAMPLES. PILIPALPINAE - Techmessa concolor Bates,
1874 (Fig. 224), Exocalopus pectinatus Broun, 1893 (Fig.
SELECTED REFERENCES. Arnett (1950), Hudson 225).
(1975), Rozen (1960).
REMARKS. The New Zealand Pyrochroidae as defined
by Pollock (1995) are equivalent to the subfamily Pili-
[71] Family PYROCHROIDAE palpinae of Pythidae proposed by Watt (1987). These
Fig. 224, 225 authors discuss the relationships between genera and the
status of the family, and provide keys to species (adults
Length 5-10 mm Tarsal formula 5-5-4 and larvae) and genera. We have followed the family
DIAGNOSIS. Body soft, flattened dorsoventrally, concept proposed by Pollock (1995), who studied the
elongate, subparallel, with elytra often slightly wid- classification, phylogeny, and geographic history of pili-
ened posteriorly from midlength, slightly glossy or matt, palpine genera worldwide, and extended the family limits
variously punctate, with decumbent and erect hairs; of Pyrochroidae.
coloration uniformly brown or black, or bicoloured with Suggested colloquial name: cardinal beetles.
pronotum orange and remainder of body dark. Head ent-
irely visible from above, gradually narrowed behind SELECTED REFERENCES. Pollock (1994, 1995), Watt
eyes (never abruptly narrowed posteriorly as in Scrap- (1987).
tiidae, Fig. 233); clypeus with an anterior membranous
area; frontoclypeal suture slightly depressed to dis-
tinctly carinate; antennal insertions exposed. Maxillary [72] Family SALPINGIDAE
palps with terminal segment expanded and truncate (incl. INOPEPLINAE)
apically. Antennae usually sexually dimorphic (males Fig. 226, 227
with increased pectination), filiform, serrate, or pec-
Length 2-10 mm Tarsal formula 5-5-4
tinate. Eyes large, projecting, entire or slightly emar-
ginate (Techmessodes). Pronotum small, two-thirds of Members of this family are represented by two distinct

—62—
forms: smalI carabid-like species, moderately convex to [73] Family ANTHICIDAE
slightly flattened, with a distinct `waist' between pro- Fig. 228-231
thorax and elytra, and elytra narrowly oval (Aegialitinae
and Salpinginae); and a staphylinid-like species of Ino- Length 2-7 mm Tarsal formula 5-5-4
peplinae with short elytra, exposed abdomen, and strongly DIAGNOSIS. Body antlike, elongate, convex, with
flattened body. elytra much broader than head and pronotum, vari-
ously punctate, clothed with decumbent and erect hairs;
DIAGNOSIS. Body usually coarsely punctate, sub- coloration black, brown, rust brown, or bicoloured with
glabrous or with long, protruding setae (Tricho- dark or yellowish spots on elytra which sometimes form
colposinus), uniformly black, dark brown, reddish brown, broad bands. Head as broad as pronotum or slightly
or bicoloured reddish brown with brown elytral spots. broader, abruptly constricted basally and forming a
Head prognathous, never abruptly narrowed behind narrow neck (exception: Lagrioida, Lagrioidinae); ant-
eyes; antennal insertions exposed. Maxillary palps ennal insertions exposed. Maxillary palps with last
with terminal segment enlarged, broad and truncate segment usually securiform. Antennae 11-segmented,
apically or narrowly elongate (Inopeplinae). Antennae filiform, usually incrassate apically but without a dis-
11-segmented, filiform, without a club (Inopeplinae) or tinct club (a weak 3-segmented club present in Macratria
with a loosely formed 3-, 4-, or indistinctly 5-segmented and Lagrioida). Eyes small to large, slightly protruding
club (Trichocolposinus). Pronotum narrowed at base, laterally. Pronotum approximately oval or quadrate in
broadened apically. Prosternum long in front of coxae. outline, widest at front and constricted (slightly so in
Procoxae globose, narrowly separated, open behind; Lagrioida) at or near base (Cotes); lateral pronotal
mesocoxae globose, and metacoxae transverse; coxae carinae absent. Elytra elongate oblong to elongate or
narrowly or moderately broadly separated (Aegialitinae). s u bparallel (some Anthicus), much broader than prono-
Abdomen with 5 ventrites, the 2 basal ones connate. tum or head; punctation confused; epipleura incomplete.
Procoxae projecting, their cavities confluent, open behind
SYNOPSIS. There are 22 native nominal species of (but closed internally); metacoxae almost touching (Mac-
salpingid in New Zealand (Watt l982a); 13 of them are ratriinae) or separated (Anthicinae, Lagrioidinae). Tarsi
represented in NZAC. The species belong to 4 genera in 3 with penultimate segment lobed. Abdomen with 5
subfamilies — AEGIALITINAE, INOPEPLINAE, and SALPINGINAE. ventrites, the basalmost 2 connate in Lagrioida.
Kuschel (1990) listed 8 species of Salpingus from Lyn-
field, Auckland. SYNOPSIS. In New Zealand there are 26 species of
Species of Salpingus are found on foliage of various anthicid, 9 of them adventive, in 7 genera and 4 sub-
trees, shrubs, and herbaceous plants, with some in sooty families — ANTHICINAE, LAGRIOIDINAE, LEMODINAE, and
moulds, and are easily collected by beating or sweeping. MACRATRIINAE (Werner & Chandler 1995). Two genera
Some larvae of Salpingus live under the bark of dead are endemic to New Zealand. Kuschel (1990) recorded 12
twigs. Species of Diagrypnodes live under bark of dead species in 4 genera from Lynfield, Auckland.
trees, including Nothofagus. The most specialised species Anthicids are general scavengers, and occur in decaying
are in Antarcticodomus, living under stones in the inter- vegetation such as compost or heaps of prunings, on
tidal zone on many offshore islands (Watt 1962, 1982b). vegetation near streams, in forest litter, and under logs.
They have an elongate terminal tarsomere with strong They may be collected by sweeping vegetation, sifting
claws, as in species of Dryopidae and Elmidae [21, 22] — organic matter, or from pitfall traps. Lagrioida brouni,
an adaptation for life on rocks in the intertidal zone. which occurs on beaches of the North Island, has related
species in Australia and Chile (Lawrence & Britton 1994).
RANGE. North ®, South ®, Stewart ®
Kermadecs ®. Three Kings ®, Chathams ®, Snares RANGE. North ®. South ®, Stewart ®
Aucklands ®, Campbell ®, Antipodes ®, Bounties ® Kermadecs O, Three Kings ®, Chathams ®, Snares O
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLES. SALPINGINAE - Salpingus bilunatus Pascoe,
1876 (Fig. 226); INOPEPLINAE - Diagrypnodes wakefieldi EXAMPLES. ANTHICINAE - Anthicus hesperi King, 1869
Waterhouse, 1876 (Fig. 227). (Fig. 228); LAGRIOIDINAE - Lagrioida brouni Pascoe, 1876
(Fig. 229); LEMODINAE - Cotes crispi (Broun, 1880) (Fig.
REMARKS. Salpingidae are poorly known, and the 230); MACRATRIINAE - Macratria exilis Pascoe, 1877 (Fig.
family is in need of taxonomic revision. 231).
Suggested colloquial name: bark mould beetles.
REMARKS. New Zealand's anthicids have been revised
SELECTED REFERENCES. Spilman (1954, 1967), and described by Werner & Chandler (1995).
Young (1991a, b). Suggested colloquial name: ant beetles.

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SELECTED REFERENCES. Abdullah (1969), Kitayama SELECTED REFERENCES. Hayashi (1972), Young
(1982), Werner & Chandler (1995), Young (1991c). (1991d).

[74] Family ADERIDAE [75] Family SCRAPTIIDAE

(= EUGLENIDAE = XYLOPHILIDAE) Fig. 233
Fig. 232
Length 1-3 mm Tarsal formula 5-5-4
Length 1-3 mm Tarsal formula 5-5-4
DIAGNOSIS. Body soft, slightly flattened, oblong to
DIAGNOSIS. Body soft, elongate, brown or black, often narrowly elongate, black, brown, or yellowish brown,
with bicoloured elytra, with head and pronotum marked- sometimes with dark spots on elytra, uniformly pubescent.
ly narrower than elytra, convex, punctate, and with Head strongly deflexed, abruptly constricted behind
erect and/or decumbent hairs. Head deflexed, abruptly eyes into a narrow neck; frontoclypeal suture distinct;
constricted behind eyes. Maxillary palps with last antennal insertions exposed. Maxillary palp with
segment securiform. Antennae 11-segmented, filiform. terminal segment securiform. Antennae filiform,
Eyes large, coarsely faceted. Pronotum small, subquad- without a club. Eyes emarginate, C-shaped, coarsely
rate, without lateral carinae. Elytra broad, oval, with faceted. Pronotum transverse, with lateral margins
distinct shoulders but lacking epipleura; punctation incomplete anteriorly; disc sometimes with 2 small basal
confused. Procoxae projecting, their cavities open; impressions. Elytra as broad as pronotum at base, with
metacoxae separate. Tarsi with penultimate segment confused punctation; epipleura incomplete. Procoxae
reduced and the preceding ones lobed. Abdomen with 2 projecting, their cavities open behind. Tarsi with
basal sternites connate or solidly fused. penultimate segments lobed beneath. Tibial spurs well
developed (Fig. 85), pubescent, not serrate. Abdominal
SYNOPSIS. Watt (1982a) reported 9 native species of ventrites free.
aderid for New Zealand. Fifteen species (many undes-
cribed) are represented in NZAC, almost all of them SYNOPSIS. There are 4 nominal species of scraptiid
belonging to a genus at present recognised as `Xylophilus.' recorded from New Zealand (Watt 1987), plus several
The assignment of New Zealand species to this European undescribed in NZAC. The described species belong to 2
genus is, however, questionable. Scraptogetus (Metasclera) genera in subfamily SCRAPTIINAE. Kuschel (1990) reported
is also recorded from New Zealand (Hudson 1934), but its 3 species of Nothotelus from Lynfield, Auckland.
assignment to Aderidae is only partially confirmed (Watt Adults occur on foliage and flowers in mainly forested
1987). Kuschel (1990) recorded 5 species of `Xylophilus' areas, and larvae are known from leaf litter.Young (1991e)
from Lynfield, Auckland. recorded scraptiid larvae from beneath bark, among
Adult aderids are commonly found on foliage, and decaying woody fibres of dead logs, and from lichens. In
larvae live in rotten wood, under bark, and in leaf litter. Australia, larvae occur under bark, in rotten wood, or in
Adults may be collected by beating foliage, in Malaise leaf litter (Lawrence & Britton 1994).
traps, or by light trapping. The larvae are probably asso-
ciated with decaying wood (Young 1991d). RANGE. North ®, South ®, Stewart
RANGE. North ®, South ®, Stewart ® Kermadecs O, Three Kings O, Chathams ®, Snares O
Kermadecs O, Three Kings ®, Chathams ®, Snares O Aucklands O, Campbell O, Antipodes O, Bounties O
Aucklands O, Campbell O, Antipodes O, Bounties O
EXAMPLE. SCRAPTIINAE - Nothotelus usitatus (Broun,
EXAMPLE. `Xylophilus' nitidus (Broun, 1893) (Fig. 1880) (Fig. 233).
232).
REMARKS. Scraptiidae may be readily distinguished
REMARKS. Aderids are similar in appearance to Anthi- from Anthicidae [73] and Aderidae [74] by the pronotum
cidae [73], but are distinguished by having the penultimate having lateral carinae and being equal in width to the
tarsal segment vestigial (cf. well developed and lobed) and elytral base, and from Melandryidae [62] by having the
the preceding segment lobed, larger and coarsely faceted eyes deeply emarginate, head strongly constricted post-
eyes, no elytral epipleura (cf. present but incomplete), and eriorly, and tibial spurs not serrate (Fig. 85).
first 2 ventrites connate or solidly fused. This group is very Suggested colloquial name: soft leaping beetles.
poorly known in New Zealand.
Suggested colloquial names: puppet beetles, ant-like SELECTED REFERENCES. Franciscolo (1964), Watt
leaf beetles. (1987), Young (1991e).

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Superfamily CHRYSOMELOIDEA (PHYTOPHAGA) alus tristis. The majority of our species are cerambycines,
Adults are characterised by having pseudotetramerous of which 6 or so species of Zorion are perhaps the most
tarsi (5-5-5-segmented with the 4th segment reduced), the strikingly colourful beetles in our fauna. The species of
head usually not prolonged into a rostrum, antennae Lamiinae were treated by Breuning (1962), who also
filiform, not clubbed, and male genitalia of a reduced provided keys for their identification.
cucujoid type (Lawrence & Britton 1994). Larvae lack a Adults feed on foliage, flowers, or bark, and some adults
distinct mandibular mola, gular area, and hypopharyngeal feed in live shoots before sexual maturation; larvae feed
bracon. All species are phytophagous. internally on bark or woody material of trees and shrubs, A
few larvae of exotic species are known to develop in seed
pods, or in stems of herbaceous plants (Booth et al. 1990,
[76] Family CERAMBYCIDAE Lawrence & Britton 1994). Many species, particularly
adventive ones, are of economic importance as pests of
[assisted by G. Kuschel]
commercial timber.
Fig. 234-237 Collected by netting, sweeping foliage and flowers, at
light, or by rearing larvae from wood.
Length 2.5-50 mm Tarsal formula 5-5-5,
appearing 4 4 4
RANGE. North ®, South ®, Stewart
DIAGNOSIS. Body elongate, subcylindrical or flattened Kermadecs ® , Three Kings ®, Chathams ®, Snares O
and wide, often broad-shouldered, pubescent or sub- Aucklands O, Campbell O, Antipodes ®, Bounties O
glabrous; coloration variable through brown, black, or
green, or of complex pattern. Head variable in form, EXAMPLES. PRIONINAE - Prionoplus reticularis White,
from prognathous with strongly projecting mandibles 1846 (Fig. 234); CERAMBYCINAE - Zorion sp. (Fig. 235),
to hypognathous or slightly opisthognathous; antennal
Oemona hirta (Fabricius,1775) (Fig. 236); LΑΜΙINΑΕ –
insertions on 2 pronounced tubercles or swellings on Xylotoles costipennis (Breuning, 1982) (Fig. 237).
frons, either between eyes and mandibles (Prioninae,
Spondylidinae) or between eyes (Cerambycinae, Lami-
inae). Maxillary palps with last segment tapering apically REMARKS. Important references for identification are
or truncate (Cerambycinae). Antennae capable of being Duffy (1953) for British species and imported timber
flexed backwards against body, elongate, extending pests, and Duffy (1963, 1968) for Australian and Oriental
beyond middle of elytra, often equal to body length or larvae. For world references on economically important
much longer. Eyes usually emarginate. Pronotum cerambycids, see Booth et al. (1990). The subfamily clas-
sification was discussed by Napp (1994) and Lawrence &
cylindrical, with lateral margin distinct, at least basally
(Prioninae), to completely absent. Mesonotum usually Newton (1995).
with stridulatory files. Elytra variable, from narrow and Suggested colloquial name: longhorn beetles.
subcylindrical to broad and flattened (Prioninae), vari-
ously sculptured. Procoxae from globular to transverse, KEY TO SUBFAMILIES OF CERAMBYCIDAE
their cavities open or closed. Tarsi with penultimate 1 Antennal insertions distant from base of mandibles in
segment reduced. Abdomen usually with 5 ventrites. lateral view ... 2
Aedeagal apodemes free, or largly so; internal sac long, —Antennal insertions close to base of mandibles (Fig.
with pigmentation confined to struts (paraprocts); styli 234), nearly in contact in lateral view ... 3
usually large. Larvae with ambulatory ampullae present on
abdominal segments 1-6 or 7. 2(1) Protibiae not grooved before apex; head weakly pro-
gnathous; maxillary palps with apical segment slightly
SYNOPSIS. One of the largest beetle families in New expanded and truncate at apex; antennae moderately to
Zealand. Hudson (1934) recorded 250 native and 6 ad- extremely elongate ... (Fig. 235, 236) .. CERAMBYCINAE
ventive species of cerambycid, whereas Watt (1982a) —Protibiae obliquely grooved on internal face before
estimated 180 valid nominal native species and 8 adven- apex; head usually hypognathous or opisthognathous;
tive. Approximately 196 native and 8 adventive species maxillary palps with apical segment fusiform; anten-
are represented in NZAC, belonging to some 56 genera (8 nae usually moderately elongate
of them exotic) in 4 subfamilies – PRIONINAE, SPONDYL- ... (Fig. 237) .. LAMIINAE
IDINAE, CERAMBYCINAE, and LAMIINAE. Kuschel (1990)
recorded 56 species in 33 genera from Lynfield, Auckland. 3(1) Pronotum with lateral carinae, at least in basal half,
The Prioninae are represented by an endemic species, and bearing spines; procoxae strongly transverse; lab-
Prionoplus reticularis, which is one of the largest beetles rum firmly united to clypeus ... (Fig. 234) .. PRIONINAE
in our fauna, reaching 50 mm in length. The Spondylidinae —Pronotum lacking lateral carinae; procoxae rounded;
are represented by an adventive Palearctic species, Arhop- labrum free ... SPONDYLΙDΙΝΑΕ

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SELECTED REFERENCES. Bain (1976a, b), Blair broad beak, notched eyes, enlarged hind femora usually
(1937), Breuning (1962), Dumbleton (1957), Hosking bearing teeth, and shortened elytra exposing the pygidium.
(1978a–e, h), Napp (1994), Watt (1983c), Zondag & Bain There are 25 native species in 4 genera of Eumolpinae in
(1976). our fauna. For a key to genera and descriptions of some
species see Shaw (1957a, b).
Our 4-9 species in 2 genera of Cryptocephalinae are
[77] Family CHRYSOMELIDAE distinct in their partially glabrous, compact, subcylindrical
(including BRUCHINAE) or cylindrical body form, hypognathous head deeply in-
[assisted by L. LeSage] serted into the prothorax, broadly separated long antennae,
and often exposed pygidium.
Fig. 238-243 Our Galerucinae include some 76 species in 2 tribes,
Length 1-10 mm Tarsal formula 5-5-5, appearing 4-4-4 Galerucini and Alticini (the latter is often given subfamily
status). Galerucini, represented by some 57 species in 3
DIAGNOSIS. Body form very diverse, elongate and endemic genera, are characterised by their prognathous or
subparallel, oval, hemispherical, cylindrical or sub- hypognathous head, with usually filiform antennae inser-
cylindrical, flattened or strongly convex, usually sub- ted close together on the frons, pronotum with distinct
glabrous but sometimes pubescent or bearing scales lateral margins, and metafemora swollen not much more
(Bruchinae, some Eumolpinae), glossy or dull (e.g., some than the mesofemora.
Galerucinae); coloration brown, black, yellowish, or mul- The Alticini, commonly called flea-beetles because of
ticolored with spots or stripes, often with a metallic shine their usually small size (1-5 mm) and ability to jump, are
(e.g., Chrysomelinae). Head prognathous, hypognathous, represented by some 19 species (6 exotic, 13 native) in 10
or opisthognathous, sometimes deeply inserted into genera. They vary in form from elongate to oval and
prothorax, sometimes bearing grooves or prominences, convex, and usually are glabrous and variable in colour—
sometimes rostrate but with rostrum no longer than wide metallic, brownish, yellowish, black, blue, or dark with
(Bruchinae); antennal insertions either close together pale stripes. They are distinct in having usually filiform
on from or welI separated, never on a prominence. antennae with narrowly separated insertions, pronotum
Antennae not capable of being flexed backwards with distinct lateral margins, and strongly enlarged meta-
against the body, usually shorter than body, filiform, femora containing the endoskeletal jumping organ.
sometimes broadening apically. Eyes usually entire. Our Chrysomelinae comprise some 41 species (includ-
Pronotum narrower than elytra or as broad as elytral base, ing 5 exotic) in 9 genera. They are usually of medium size,
with or without lateral margins. Mesonotum lacking predominantly glossy and metallic, with body shape var-
stridulatory files. Elytra entire or shortened and leaving iable from narrowly elongate to broad or hemispherical
pygidium exposed, striate or with confused punctures .Tarsi and of differing convexity, and head usually oblique, with
with 4th segment usually short and concealed at base of front margin of clypeus approximately truncate, antennae
preceding segment. Abdomen with 5 ventrites; basal filiform and widely separated, and procoxae transverse.
ventrites not fused. Aedeagal apodemes fused to form a The majority of chrysomelids feed on leaves, though
spoon-like roof over internal sac, which is short and in some are known to consume pollen and anthers. Bruchines
dorsal view completely covered by fused apodemes; feed mainly on legume seeds, and all but one are con-
ovipositor with proximal hemisternites short, weakly sidered to be agricultural pests. In New Zealand chryso-
pigmented or membranous; styli usually very small or melid adults are commonly collected from low-growing
absent. Larvae without ambulatory ampullae on abdom- plants or bushes. Many species are host-specific. The fam-
inal segments. ily contains a relatively large number of pests of cultivated
plants, some of them transmitting plant viruses. Paropsis
SYNOPSIS. Hudson (1934) recorded 155 native species charybdis (Chrysomelinae), self-introduced to New Zea-
of chrysomelid and 1 adventive species of bruchid (here as land from Australia, is causing severe damage in gum
Bruchinae). Watt (1982a) recorded 146 native species and plantations (Selman 1963, Bain 1977a). For cosmopolitan
7 adventive or introduced, including 1 bruchine. Five species of economic importance see Booth et al. (1990).
more introduced bruchines are now represented in NZAC, Larvae have very diverse feeding habits, foraging exter-
making a total of some 134 native species and 19 adventive nally or internally on leaves, or stems, or roots. In general,
or introduced. These belong to some 33 genera in 5 sub- larvae of Cryptocephalinae feed on dead leaves accumu-
families – BRUCHINAE, CRYPTOCEPHALINAE, EUMOLPINAE, lated on the ground (known only from exotic species),
CHRYSOMELINAE, and GALERUCINAE. Kuschel (1990) repor- those of Eumolpinae occur in the soil and probably are
ted 20 species in 12 genera from Lynfield, Auckland. root feeders, those of Chrysomelinae feed on foliage, and
The Bruchinae, represented by 6 species in 4 genera, are many Galerucinae are miners of roots and leaves, though
distinct in having an ovate body, usually broad posteriorly, some feed externally on foliage, fruits, and flowers; all our
the head concealed from above and prolonged into a short, introduced Bruchinae feed on legume seeds.

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 Several beneficial leaf beetles have been deliberately —Body oval to rounded or robust, but not cylindrical
introduced to New Zealand for weed control: Alticini — (Fig. 239, 243), often strongly convex and constricted
Agasicles hygrophila, originally from South America, for anteriorly; antennae usually short ... 5
control of alligator weed (Alternanthera philoxeroides),
and Longitarsus jacobaeae, from Europe, for control of 5(4) Head oblique, with frontal margin straight or broadly
ragwort (Senecio jacobaea); Chrysomelinae — Chrysolina arcuate; clypeus distinct, more or less membranous;
hyperici and C. quadrigemina, from Europe, for control of body usually broadly oval, robust, slightly depressed,
St John's wort (Hypericum); Bruchinae — Bruchidius vil- glabrous; pygidium usually not exposed; 3rd tarsal
losus, from the United Kingdom, for biological control of segment usually bilobed ... (Fig. 239) .. CHRYSOMELINAE
broom (Cytisus scoparius) (see Syrett et al. 1996). —Head hypognathous, with frontal margin usually
broadly emarginate; clypeus indistinct; body elongate-
RANGE. North ®, South ®, Stewart ® robust to oval and convex, sometimes pubescent, with
Kermadecs ®. Three Kings ®, Chathams ®, Snares Ο elytra often broader than pronotum; pygidium usually
Aucklands O, Campbell O, Antipodes O, Bounties O at least partly exposed; 3rd tarsal segment distinctly
bilobed ... (Fig. 243) .. EUMOLPINAE
EXAMPLES. BRUCHINAE—Bruchidius villosus (Fabricius,
1792) (Fig. 238); CHRYSOMELINAE - Chrysolina hyperici SELECTED REFERENCES. Bain (1977a), Borowiec
Forster, 1771 (Fig. 239); GALERUCINAE —Agasicles hygro- (1987), Jolivet & Cox (1996), Kay (1980), Lawson
phila Selman & Vogt, 1971 (Fig. 240), Adoxia vulgaris (1991), Pfaffenberger (1991), Reid (1995), Seeno &
(Broun, 1880) (Fig. 241); CRYPTOCEPHALINAE - Arnomus Wilcox (1982), Selman (1963), Shaw (1957a, b).
brouni Sharp, 1876 (Fig. 242); EUMOLPINAE - Eucolaspis
brunnea (Fabricius, 1792) (Fig. 243).
Superfamily CURCULIONOIDEA
REMARKS. New Zealand's chrysomelids are poorly (RHYNCHOPHORA)
known and badly in need of taxonomic revision. Many of [assisted by G. Kuschel and B.M. May]
our genera which are now considered as native may reveal
a broader origin in the light of detailed studies of the entire Adult weevils are distinct in having the anterior part of the
Australasian fauna. head extended into a long or short rostrum (Fig. 6, 7), in
Suggested colloquial name: leaf beetles (Alticini — flea- which the mouthparts are often strongly modified (fusion
beetles). of labrum, fusion of lacinia and galea, usually reduction of
mandibles, formation of rigid palps). Additional diagnos-
KEY TO SUBFAMILIES ΟF CHRYSOMELIDAE tic characters are antennae not filiform, usually clubbed,
1 Body ovate, slightly broadening posteriorly, bearing with a long scape sometimes fitting into grooves on either
recumbent setae or scales; head produced anteriorly side of rostrum (Fig. 7), body rigid, with closed procoxal
and mesocoxal cavities, procoxae globular with concealed
into a short, broad rostrum, strongly deflexed, con-
trochantins, 2 or more abdominal sternites connate, and
stricted behind eyes into a distinct neck
tarsi pseudotetramerous (5-5-5-segmented but appearing
... (Fig. 238) .. BRUCHINAE
—Body not ovate, lacking setae and scales (exception: 4-4-4-segmented).
some Eumolpinae); head not rostrate, usually not Larvae are distinct in their cylindrical body form (Fig.
99), with abdominal segments 2-7 unsclerotised and with
strongly deflexed, without a neck, and usually deeply
... 2 dorsal plicae or folds, the presence of a hypopharyngeal
inserted into prothorax
bracon (except in some leaf miners), maxillae with lacinia
2(1) Antennae closely adjacent, inserted between eyes on and galea united to form a mala, and legs absent or ves-
frons (Fig. 240) ... 3 tigial (May 1993).
—Antennae well separated, inserted above mandibular
bases (Fig. 239, 243) ... 4
[78] Family NEMONYCHIDAE
3(2) Metafemur moderately to strongly enlarged, in
Alticini adapted for jumping, with an endosclerite, in [assisted by G. Kuschel]
Galerucini not so ... (Fig. 240, 241) .. GALERUCINAE Fig. 244
—Metafemur not enlarged, not adapted for jumping,
Length 2.0-2.8 mm Tarsal formula 5-5-5,
without an endosclerite ... 5 appearing 4-4-4
4(2) Body cylindrical or subcylindrical, with pronotum DIAGNOSIS. Body moderately elongate, narrowly
about as wide as elytral base; antennae usually long oval in outline, rust brown to dark brown, with yellowish-
and filiform; pygidium usually partly exposed white decumbent pubescence and some scattered erect
... (Fig. 242) .. CRYPTOCEPHALINAE and inconspicuous setae. Head abruptly constricted in

— 67 —
front of eyes, forming an elongate, narrow rostrum, hairs or scales forming patterns. Head produced
somewhat flattened apically, slightly shorter than pro- anteriorly into a rostrum lacking a paired gular suture
thorax in male and longer in female; antennal insertions ventrally and usually short, broad, and flattened (not
in apical third of rostrum, distant from mandibular cylindrical), rarely moderately elongate. Labrum distinct,
sockets by at least twice width of scape. Labrum free, free, separated by a groove from rest of head. Mandibles
small, sinuously triangular, with 4 short peg-like setae well developed. Maxillae with long, flexible palps and a
at apex and 2 pairs of setiferous punctures dorsally. distinct lacinia. Antennae straight, not elbowed (ex-
Mandibles acute. Maxillary palps short, flexible, loosely ception: slightly elbowed in males of Hoherius), in some
articulated. Antennae straight, with scape short, ap- males much longer than body length, with a more or less
proximately as long as 2 following segments combined, distinct 2- or 3-segmented club, and with scape received
and with a loose, 3-segmented club. Eyes round lat- into a scrobe on side of head. Pronotum usually with a
erally, coarsely faceted. Pronotum not carinate. Elytra transverse sub-basal carina joined to lateral carinae of
distinctly and regularly striate. Hind wings well dev- varying length. Elytra internally with a supracostal
eloped. Procoxae conical, their cavities closed. Tibiae flange, and usually with a scutellary striole. Procoxae
with 2 spurs in both sexes. Tarsal claws with a denti- globose, projecting, their cavities closed behind. Tibiae
form appendage. Abdominal ventrites freely articulated; lacking spurs. Abdomen with 5 ventrites, the basal 4
pygidium concealed by elytra. fused together; pygidium exposed beyond elytra.

SYNOPSIS. New Zealand has 4 species of nemonychid, SYNOPSIS. There are 58 endemic New Zealand species
of which 1 native species is described and 3 are undes- of anthribid, and 3 introduced (2 adventive, 1 commonly
cribed (G. Kuschel, pers. comm.); all belong to the sub- intercepted), belonging to 28 genera in 2 subfamilies —
family RHINORHYNCHINAE. ANTHRIBINAE and CHORAGINAE. Of the endemic species, 5
Rhinorhynchus rufulus represents an archaic lineage are confined to the Three Kings, 6 have been recorded only
associated with Podocarpaceae (Kuschel 1995). Adults from the North I. and 7 from the South I., 6 are confined to
occur in epiphytes of podocarp forests and occasionally in the Chathams, 1 is found only on Bounty I., 1 is shared by
forest litter; a few have been collected in Northland from Stewart I. and The Snares, and 1 is present on Stewart I.,
logs of kauri (Agathis australis). Kuschel (1990) recorded The Snares, and the Auckland Is. The remainder are more
this species from Lynfield, Auckland, on tanekaha (Phyl- widely distributed on the main islands. The majority of
locladus trichomanoides), miro (Prumnopitys ferruginea), endemic species restricted to a particular zoogeographic
and kahikatea (Dacrycarpus dacrydioides). Larvae, desc- region are flightless.
ribed and illustrated by May (1993), are pollen feeders Most of our anthribids are associated with standing
developing in male cones (strobili) of Podocarpaceae. vegetation in natural plant communities. Holloway (1982)
established the following groups of habitats for adults: (1)
RANGE. North ®, South ®, Stewart exclusively on standing vegetation; (2) on standing vege-
Kermadecs O, Three Kings O, Chathams O, Snares O tation or in litter; (3) exclusively in litter; and (4) in or on
Aucklands O, Campbell O, Antipodes O, Bounties O littoral lichens or fungi. Adult anthribids feed on fungi
(mainly Ascomycetes and Fungi Imperfecti) and lichens
EXAMPLE. RHINORHYNCHINAE — Rhinorhynchus rufulus (Holloway 1982, Kuschel 1995). The fungal material con-
(Broun, 1880) (Fig. 244). sists of spores, fragments of hyphae, and ruptured fruiting
bodies. Four species feed on hyphae and ascospores of
REMARKS. Suggested colloquial name: straight-horned sooty moulds, which grow on the honeydew deposited by
weevils. some scale insects (Coccoidea). Larvae are mostly endo-
phytic in dead and dying branches of trees and shrubs
SELECTED REFERENCES. Kuschel (1983, 1995), May (Holloway 1982).
(1993).
RANGE. North ®, South ®, Stewart
Kermadecs O, Three Kings ®, Chathams ®, Snares
[79] Family ANTHRIBIDAE Aucklands ®, Campbell O, Antipodes O, Bounties
[assisted by B.Α. Holloway]
Fig. 245, 246 EXAMPLES. ANTHRIBINAE Sharpius venustus (Broun,
—

1914) (Fig. 245); CHORAGINAE Dysnocryptus pallidus

—
Length 0.8-7 mm Tarsal formula 5-5-5, Broun, 1893 (Fig. 246).
appearing 4-4-4
DIAGNOSIS. Body moderately elongate to ovate, REMARKS. For keys to the genera and species occurring
moderately to strongly convex, usually strongly sclero- in New Zealand see Holloway (1982).
tised, clothed with black, brown, greenish, and/or white Suggested colloquial name: fungus weevils.

— 68 —
KEY TO SUBFAMILIES OF ANTHRIBIDAE for oviposition and larval development, pupation occur-
[Slightly modified from Holloway 1982] ring in situ (May 1993). Adults of Pachyurinus are often
beaten from Podocarpaceae, but the host trees of known
First and 2nd antennal segments not arched, usually larvae are angiosperms, Nothofagus and Toronia (May
symmetrical, but if asymmetrical then antennae very 1993). Aralius wollastoni lives subcortically in dying or
slender and reaching back beyond middle of elytra; dead branches of Pseudopanax arboreus and P. lessonii,
these 2 segments no more convex on external margin
where the thin bark layer has not yet become dry (Kuschel
than on internal margin when antennae folded against 1990, May 1993). For general information on natural
body; scrobes either lateral or, if dorsolateral, then 1st
history see May (1993) and Kuschel (1995).
antennal segment pyriform or obliquely truncate at
base ... (Fig. 245) .. ANTHRIBINAE RANGE. North ®, South O, Stewart O
—First and 2nd antennal segments arched and asym-
Kermadecs O, Three Kings O, Chathams O, Snares O
metrical, more convex on external margin than on Aucklands O, Campbell O, Antipodes O, Bounties O
internal margin when antennae folded against body;
scrobes always dorsal ... (Fig. 246) .. CHORAGINAE EXAMPLES. BELINAE - Cyrotyphus tridens (Fabricius,
1792) (Fig. 247); AGLYCYDERINAE - Aralius wollastoni
SELECTED REFERENCES. Holloway (1971, 1982), (Sharp, 1876) (Fig. 248).
Kuschel (1995), May (1993), Zimmerman (1994).
REMARKS. Suggested colloquial name: austral weevils.
[80] Family BELIDAE KEY TO SUBFAMILIES OF BELIDAE
[assisted by G. Kuschel] OCCURRING IN NEW ZEALAND
Fig. 247, 248 [by G. Kuschel]
Length 3-13 mm Tarsal formula 5-5-5, appearing Antennae lacking a distinct club (a weak, loosely defined
4-4-4; or 4-4-4, appearing 3-3-3 4-segmented club rarely present); tarsal formula 5-5-
5, but appearing 4-4-4 because of vestigial 4th seg-
DIAGNOSIS. Body elongate and subparallel or broad-
ment; tarsal segment 1 at least as long as segments 2
ly, irregularly oval, with head and pronotum distinctly
narrower than elytra and tapering anteriorly; surface and 3 combined ... (Fig. 247) .. BELINAE
especially of elytra sometimes tuberculate, with decum- —Antennae with a distinct 2-segmented club; tarsal
formula 4 4 4, appearing 3-3-3 because of vestigial
bent or erect pubescence or scales. Head either broad,
3rd segment; tarsal segment 1 shorter than segments 2
abruptly constricted behind eyes, and with rostrum
and 3 combined ... (Fig. 248) .. AGLYCYDERINAE
short and broad (Aglycyderinae) or not constricted
behind eyes and with rostrum elongate and usually
SELECTED REFERENCES. Kuschel (1995), May
broad (Belinae); antennal insertions usually in basal
(1993), Zimmerman (1994).
half of rostrum. Labrum not visible. Antennae straight,
with a well defined 2-segmented club (Aglycyderinae) or
without a club, rarely with a weak and loosely defined
[81] Family BRENTIDAE
4-segmented club (Belinae). Pronotum with sides sub-
parallel or converging apically, lacking lateral margins. (incl. ΑΡΙΟΝΙΝΑΕ)
Elytra variable in shape, often subparallel in part, some- [assisted by G. Kuschel]
times with protuberances (Belinae) or with coarsely punc- Fig. 249, 250
tate striae and projecting short hairs and scales. Procoxae Length 2-4//18-75 mm

Tarsal formula 5-5-5,
projecting, their cavities broadly closed behind but some- appearing 4-4-4
times with narrow lateral extensions. Tarsi with 4th or
3rd segment reduced. Abdomen with 5 ventrites; DIAGNOSIS. Body either extremely narrowly elongate,
pygidium concealed. with head as long as pronotum and elytra combined or
slightly shorter in females (Brentinae), or moderately
SYNOPSIS. Belids are represented in New Zealand by 2 elongate and pyriform, with head as long as elytra or
subfamilies – BELINAE with 10 described native species in shorter (Apioninae), subglabrous or pubescent. Head
2 genera, and AGLYCYDERINAE with 1 endemic New with rostrum long and narrow; gular suture distinct
Zealand species. Kuschel (1990) recorded the latter from and merged into one; antennal insertions at base, at apex,
Lynfield, Auckland, and May (1993) from Whangamoa or in middle of rostrum. Labrum not visible. Maxillae
and Cuvier Island. and labium reduced; maxillary palps rigid and labial
Adults of Cyrotyphus are usually found in association palps reduced. Antennae straight or (Neocyba) slightly
with Podocarpaceae, but utilise dead wood of many kinds geniculate, lacking a club (Brentinae) or club present

-69-
(Apioninae). Prothorax without lateral carinae. Elytra DIAGNOSIS. Body highly variable in form, robust,
concealing pygidium. Procoxae projecting, their cav- strongly sclerotised, convex, subglabrous or with scales
ities closed. Protibiae without a grooming device on and/or bristles. Head with a long or short rostrum.
inner face (consisting of dense vestiture in a groove or Labrum absent. Maxillae reduced, with short rigid
impression). Tarsi with penultimate segments bilobed. palps; gular sutures fused. Antennae geniculate, with a
Abdomen with first 2 visible sternites fused together, long scape and more or less compact club (Fig. 7, 28).
much longer than remaining sternites. Procoxae projecting, their cavities closed. Tarsi with
penultimate segment minute and concealed at base of
SYNOPSIS. New Zealand's brentid fauna comprises 3 lobed 3rd segment (exception: Platypodinae).
genera, with 1 very distinctive native species in subfamily
BRENTINAE and 5 species (1 exotic, 2 undescribed) in sub- SYNOPSIS. Curculionids are the most diversified group
family APIONINAE. of beetles, with some 50 000 species in the world fauna
The giraffe weevil, Lasiorhynchus barbicornis, occurs (Lyal 1993). They are also the largest beetle family in New
on a variety of trees including kauri (Agathis australis), Zealand. Watt (1982a) recorded 1279 native and 42 ad-
pigeonwood (Hedycarya arborea), rewarewa (Knightia ventive species. There are 1496 native and 46 adventive
excelsa), tawa (Beilschmiedia tawa), and karaka (Coryno- species in 231 genera represented in NZAC, belonging to
carpus laevigatus). The female chews a cavity in the bark 6 subfamilies in the new system of classification proposed
of dying or suppressed trees and in freshly felled logs for by Kuschel (1995) — BRACHYCERINAE (49 genera), CURCU-
oviposition (Meads 1976, May 1993). The gorse weevil, LIONINAE (126), DRYOPHTHORINAE (Rhynchophorinae) (4),
Exapion ulicis, was introduced for biocontrol of gorse COSSONINAE (36), SCOLYTINAE (14), and PLATYPODINAE (2).
(Ulex europaeus), and Neocyba metrosideros, a native Kuschel (1990) recorded 184 species (24 adventive) in
species, occurs on pohutukawa (Metrosideros excelsa) 102 genera and 6 subfamilies from Lynfield, Auckland.
(Kuschel 1990). So far as is known, weevils are exclusively phyto-
phagous. They occur in a wide range of habitats: some feed
RANGE. North ®, South ®, Stewart O on leaves, others live in the soil and feed on roots; some
Kermadecs O, Three Kings O, Chathams O, Snares O bore into stems, and others mine between the upper and
Aucklands O, Campbell O, Antipodes O, Bounties O lower surface of leaves (May 1993). Larvae are cryptic,
and usually feed internally in plant tissue; some feed on
EXAMPLES. ΒRΕΝΤINΑΕ — Lasiorhynchus barbicornis flowers and in developing ovaries (May 1987, 1993).
(Fabricius, 1792) (Fig. 249); ΑΡΙΟΝINΑΕ — Neocyba metro- Females of Brachycerinae oviposit in the soil, and the
sideros (Broun, 1880) (Fig. 250). larvae feed on roots (May 1993). The curculionines have
many host-specific species, with larvae usually feeding
REMARKS. Suggested colloquial name: primitive wee- endophytically on living plants (May 1993). Cossonines
vils. and some rhynchophorines utilise dead tissue of bark or
rotten wood. Adults of Scolytinae bore through bark to
KEY TO SUBFAMILIES OF BRENTIDAE construct a nursery gallery, later expanded by the larvae.
Body extremely elongate and narrow, length 18-75 mm; Platypodinae, known as pinhole borers, cause damage to
head as long as remainder of body, or nearly so; timber by their tunnelling activities and by `sap staining'
antennae without a club; tibial spur present; trochan- from secondary fungal infection, making them considerable
ters not elongate, hence femora contiguous with coxae pests in the timber industry. Both scolytines and platy-
at base ... (Fig. 249) .. BRENTINAE podines feed on ambrosia fungi cultivated in the galleries.
—Body moderately elongate and pyriform, length less Several weevil species have been accidentally intro-
than 4 mm; head as long as elytra or shorter; antennae duced into Νew Zealand, and some of these are considered
with a club; tibial spur absent; trochanters elongate, to be pests of agriculture, forestry, or gardens. For a list of
hence femora distinctly disjunct from coxae at base adventive species see Kuschel (1972).
... (Fig. 250) .. ΑΡΙΟΝΙΝΑΕ
RANGE. North ®, South ®, Stewart ®
SELECTED REFERENCES. Kuschel (1995), May Kermadecs ®, Three Kings ®, Chathams ®, Snares
(1987, 1993), Zimmerman (1994). Aucklands ®, Campbell ®, Antipodes ®, Bounties O

EXAMPLES. ΒRΑCHYCERIΝΑΕ —Anagotus turbotti (Spiller,

[82] Family CURCULIONIDAE 1942) (Fig. 251), Mandalotus miricollis (Broun, 1917)
[assisted by B.M. May and G. Kuschel] (Fig. 252); CURCULIONINAE —Andracalles spurcus (Broun,
1880) (Fig. 253), Myrtonymus zelandicus Kuschel, 1990
Fig. 251-260 (Fig. 254), Stephanorhynchus lawsoni Sharp, 1876 (Fig.
255); COSSONINAE — Macroscytalus remotus (Sharp, 1878)
Length 0.7-30 mm Tarsal formula 5-5-5 or 4-4-4

— 70 —
(Fig. 256), Xenocnema spinipes Wollaston, 1873 (Fig. 4(3) Mandibular socket deep (deeply emarginate); hypo-
257); SCOLYTINAE — Hylastes ater (Paykull, 1800) (Fig. stomal tooth long, usually sharply pointed; pharyngeal
258), Phloeosinus cupressi Hopkins, 1903 (Fig. 259); process long, about as long as a mandible or longer
PLATYPODINAE- Platypus apicalis White, 1846 (Fig. 260). ... (Fίg. 256, 257) .. COSSONINAE
—Mandibular socket shallow (shallowly emarginate);
REMARKS. The most striking New Zealand weevils are hypostomal tooth absent or indistinct; pharyngeal pro-
in the brachycerine genus Anagotus, which includes both cess less than half of mandibular length, or absent ... 5
small species and the largest, often called giant weevils.
All are flightless and have specialised habitat requirements. 5(4) Procoxa antemedian; antennal funicle 5-7-segmented;
They are considered endangered because they are prone to first 2 visible sternites fused; sternite 2 distinctly lon-
predation by the Polynesian rat (kiore, Rattus exulans), ger than sternite 3; tarsi not or slightly longer than
and many are surviving on rat-free islands (see Examples tibiae; sternite 9 of male present
of Species and Appendix 1). Evidence that they were once ... (Fig. 258, 259) .. SCOLYTINAE
broadly distributed on the mainland includes fragmentary —Procoxa postmedian; antennal funicle 4-segmented;
remains found in cave deposit at Waitomo (approx. 1700 first 2 visible sternites usually free; sternite 2 not lon-
years old) and in the Pureora Buried Forest deposits ger than sternite 3; tarsi distinctly longer than tibiae;
formed during the Taupo eruption (circa A.D. 130). sternite 9 of male usually absent
New Zealand species of Cryptorhynchini (Curculio- ... (Fig. 260) .. ΡLATVPODΙNE
ninae), originally treated as a subfamily, were revised by
Lyal (1993). New Zealand species of Molytini are revised SELECTED REFERENCES. Bain (1976c, 1977b—e),
by Craw (in press), under subfamily Molytinae. Barratt (in press), Craw (in press), Hosking (1978g, 1979),
Suggested colloquial name: weevils. Kuschel (1964, 1966, 1969, 1970, 1971, 1972, 1987,
1995), Lyal (1993), Marshall (1926, 1937, 1944, 1953),
KEY TO SUBFAMILIES OF CURCULIONIDAE May (1966, 1987, 1993), Milligan (1978, 1979b, c),
OCCURRING IN NEW ZEALAND Zimmerman (1992—[98]), Zondag (1976, 1977).
[Modified from Kuschel 1995]
— v —

1 Epistome (dorso-apical part of rostrum, Fig. 6) usually

uneven, raised or impressed in relation to surrounding
area; mandibles usually multisetose; rostrum usually REFERENCES
short and thick, not sexually dimorphic; underside of Abdullah, M. 1969: The natural classification of the
elytra lacking stridulatory files; sclerolepidia absent; family Anthicidae with some ecological and etholo-
ventrite 9 of male with bladal part extensively sclero- gical observations (Coleoptera). Deutsche entomolog-
tised, the arms pigmented, broad, usually discontinuous ische Zeitschrift, MN.F. 16: 323-366.
with apodeme ... (Fig. 251, 252) .. BRACHYCERINAE
—Epistome even, not raised or impressed in relation to 1973: The systematic position of Cisidae (Hetero-
surrounding area; mandibles usually more sparsely mera) including a catalogue of the world and com-
setose; rostrum mainly long and slender, usually sex- ments on central European families of Cucujoidea
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stridulatory files; sclerolepidia frequently present; Allison, A.; Samuelson, G.A.: Miller, S.E. 1993: Patterns
ventrite 9 of male with bladal part largely mem- of beetle species diversity in New Guinea rain forest as
branous, the arms pigmented, usually narrow, fused to revealed by canopy fogging: preliminary findings.
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2(1) Mesotibia and metatibia with distal or ascending Arndt, E. 1993: Phylogenetische Untersuchungen larval-
combs ... (Fig. 253-255) .. CURCULIONINAE morphologischer Merkmale der Carabidae (Insecta:
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Oedemeridae and their type species. Journal of the
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Washington Academy of Sciences 40(7): 217-225.
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with 7 segments, if apparently fewer then because of 1973: The beetles of the United States. Michigan,
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 GLOSSARY OF TECHNICAL TERMS biodiversity, the totality of all the different plants, animals,
Most of the definitions below have been formulated with fungi, and microorganisms, their genetic components, and their
reference to Lawrence & Britton (1994), Snodgrass (1935), and ecological associations. Normally used in a narrower sense,
Nichols (1989). e.g., biodiversity of beetles from a particular area.
aciculate, slender and needlelike. bionomics, the life habits, reproduction, and adaptations of
organisms.
acone, a condition of the insect eye in which the ommatidium
has neither a crystalline nor a liquid cone, in place of which biota, the fauna and flora of a defined habitat or region.
there is a group of elongate transparent cells; eye with open bipectinate (of antennae), see p. 14.
rhabdoms. bladal, pertaining to the bladelike portion of sternite 9.
adventive (of species, populations), having arrived only cambium, an area of growing and differentiating cells in the
recently in an area under discussion, either through dispersal or stems and roots of vascular plants.
through inadvertent or deliberate human agency. campodeiform (of larvae), at least in the early stages
aedeagus, the male copulatory organ, derived from the resembling Campodea (Diplura), i.e., elongate, well
posterior portion of the ejaculatory duct and consisting of three sclerotised, dorsoventrally flattened, usually long legged,
parts: the basal tegmen, which is often composed of a basat predacious, with prognathous head and usually with terminal
piece (phallobase) and usually two lateral lobes called abdominal processes (Fig. 100).
parameres; the penis (median lobe), which is usually enclosed cantharidin, a defensive chemical found in the haemolymph
at the base by the tegmen; and the internal sac (endophallus), of adult Meloidae and Oedemeridae, blistering human skin.
which is primarily membranous but often bears internal canthus, a shelf-like lateral expansion of the head, practically
sclerites and a sclerotised flagetlum (after Lawrence & Britton
,
covering the base of the first antennal segment (e.g., in
1994). Tenebrionidae: Fig. 11, 1).
allopatric (of populations or species), occupying mutually capitate (of antennae), see p. 14 and Fig. 23, 187.
exclusive geographic areas (cf. sympatric). capsuIe, a small, closed, container-like structure; often
alluvial (of soils), deposited by the action of water. referring to insect head capsule (Fig. 1, 2, 4, 6, 7).
ambrosia, the fungus cultivated by wood-boring Scolytinae cardo (pl. cardines), the basal division of the maxilla in
and Platypodinae; more specifically, the part of the fungus that Coleoptera (Fig. 5).
grows out into the burrows and is eaten by the beetles. carina (pl., carinae), an elevated ridge or keel.
ampulla (pl. ampullae), a blisterlike structure; often referring cercus (pl. cerci), an appendage (generally paired) of abdom-
to larval structures, e.g., ambulatory ampullae inal segment 9, but often appearing to belong to abdominal
antemedian, situated before the middle (cf. postmedian). segment 10.
antenna (pl. antennae), paired segmental appendages borne on cladistic analysis, technique for grouping taxa according to
either side of head, and bearing sensory receptors (Fig. l, 17- probable ancestral relationships, based on measures of
33). similarity and difference.
antennal cleaning organ, an excavation of the protibia lined clavate (of antennae), see p. 14 and Fig. 23-29.
with a comb-like setal fringe for cleaning antennae, e.g., in clavicorn, with antennae more or less distinctly clubbed; hence
Carabidae. Clavicornia, a group of beetles with this feature.
antennomere, one of the antennal divisions (usually 11 in click mechanism, in adult Elateridae and Eucnemidae, a com-
Coleoptera), which are jointed together and form the entire plex `peg and socket' structure consisting of the long prosternal
antenna (Fig. l). process and mesosternal cavity; sudden forcing of the process
anterior pit (= anterior tentorial pit), a small round, internal out of the cavity causing convulsive movement of the
depression between clypeus and frons (see also tentorial pit) prothorax relative to the hind body, often resulting in a jump
(Fig. l). and/or an audible click (Fig. 54).
apodeme, an ingrowth of the insect exoskeleton (q.v.) to which club (of antennae), seep. 14 and Fig. 23-31.
muscles are attached. clypeus, part of the insect head capsule below the frons, to
apomorphy, a derived character or character state (cf. plesio- which the labrum is attached anteriorly (Fig. 1).
morphy); hence apomorphic. coeval (of two or more organisms), having evolved contempo-
approximated, in close physical relationship. raneously in the geological time scale.
apterous, without wings. comb, a row or patch of specialised spines, usually on tibiae
articulation, mobile joint between two rigid structures. (Fig. 162).
aspirator, suction device for capturing small insects. compound eye, in many arthropods, an aggregation of visual
Barber's fluid, beetle relaxing fluid: ethanol 95% (53 parts), elements (ommatidia) each of which corresponds with a single
water (49 parts), ethyl acetate (19 parts), and benzol (7 parts). facet of the cornea.
Berlese funnel, see p. 17. conidia, asexual spores of fungi.
biforous (of a spiracle), with two extended openings, or with connate (of stemites), immovably joined together, cf. the more
one but in appearance similar to the double spiracle. usual movable (Fig. 91) and separate configuration.

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coronal suture, the median unpaired part of the epicranial endophallus, part of the aedeagus.
suture. endopleuron (pl., endopleura), the reduced pleuron of Poly-
costate, with longitudinal raised ribs or ridges (costae), much phaga which is fused with the trochantin and hidden internally.
coarser than carinae. endosclerite, an internal sclerite, e.g., of the metafemur in
coxa (pl., coxae), the basal segment of the leg, articulating to Alticini, sometimes called a metafemoral spring, which
the body (Fig. 57-65). constitutes a skeletal jumping organ.
coxal cavity, the opening or space in which the coxa articu- epicranium, upper part of head capsule from frons so neck,
lates. May be laterally closed by sterna alone, i.e., excluding including frons, vertex, and genae (Fig. l, 6, 7). Hence
pleural elements, or laterally open, when only partly or epicranial stem, οr coronal suture, and epicranial suture, a
completely closed by pleural elements (Fig. 40-45). Y-shaped suture on the head capsule above the antennae (Fig.
crenulate (of margins), evenly notched with small, rounded 113).
teeth (Fig. 163, 164 — hind margin of pronotum). epigeal, living on the surface of the earth (cf. hypogeal).
Cretaceous, third period of the Mesozoic Era, extending about epimeron (pl. epimera), the posterior division of a thoracic
135-70 million years before the present. pleuron, delimited anteriorly by the pleural sulcus (Fig. 2, 3).
cryptic, hidden or concealed, referring to (a) life habit οr (b) epiphyte, a plant growing nonparasitically upon others,
taxonomic status, e.g., cryptic species. especially in the upper branches of trees, e.g., some ferns and
crypto-, hidden, as in, e.g, cryptopleuron. orchids.
cryptonephridism, a condition in which the Malpighian tubes epipleuron (pl., epipleura), in adult beetles, deflexed or
are closely associated with the rectum, forming a convoluted inflexed lateral portion of pronotum οr elytra (Fig. 2, 3, 34,
layer over its surface. 35); hence epipleural carina, a sharp ridge at lateral edge of
cucujoid (of the aedeagus), a configuration in which the epipleuron.
phallobase forms a ring around the penis. episternum (pl., episterna), anterior sclerite of pIeuron,
cupule (of antennae), transverse and concave segment delimited posteriorly by pleural sulcus (Fig. 2).
preceding the club (e.g., in Hydrophilidae, Fig. 27); hence eplstome, sclerite immediately behind labrum (Fig. 6, b).
cupuliform. eruciform (of a larva), resembling a caterpillar (Fig. 97).
cuticle, outer layer of the insect i ntegument. excavate(d), hollowed out, concave, as of a margin.
decumbent, bent downwards. exoskeleton, extemal skeleton of arthropods, consisting of hard
deflexed, abruptly bend downwards. or flexible cuticle to the inside of which muscles are attached.
denticulate, beset with tiny teeth or notches. explanate (of a margin), spread out and flattened (Fig. 217 —
detritivore, an animal that feeds on organic detritus. pronotal margin).
dichotomy, a branching of a single stem into two equal and exuviae, the moulted, cast-off skin of larvae or nymphs.
diverging parts; hence dichotomous key, a key with couplets femoral lines, fine ridges on basal abdominal sternite running
divided into pairs of opposing character sets. backwards and outwards from inner end of metacoxae (Fig.
dimorphic, existing in two forms; hence sexually dimorphic, 92).
differing between male and female. femur (pl., femora), third and usually stoutest segment of the
disc, medial surface of a body sclerite; e.g., central surface of leg, articulated to trochanter proximally and to tibia distally
pronotum. (Fig. 57).
disjunct, separated or lacking continuity; often referring to filiform (of antennae), see p. 14 and Fig. 17, 18.
distribution of species or populations. flabellate (of antennae), see p. 14 and Fig. 33.
distal, near or towards the free end of any appendage; that part flagellum (pl., flagella), third part of antenna beyond pedicel.
of a segment farthest from the body (cf. proximal). Also part of the aedeagus.
diurnal, active or habitually flying by day (cf. nocturnal). flight interception trap, see p. 17.
dorsal, on the functionaIly upper surface (cf. ventral). Hence fossorial (of legs), adapted for burrowing in soil.
dorsoapical, on the upper surface and towards the apex; fovea (pl., foveae), a pit; e.g., on head and pronotum of
dorsolateral, at the top and to the side; dorsoventral, in a line Pselaphinae, Staphylinidae (Fig. 126).
from the upper to lower surface. frons, upper anterior portion of head capsule, usually a distinct
dorsum, the anatomically upper surface. sclerite between epicranium and clypeus (Fig. l).
ectoparasite, an external parasite (cf. endoparasite). frontoclypeal suture, suture between frons and ctypeus (Fig.
elytron (pl., elytra), the leathery forewing of beetles, serving as l).
a covering for the hind wings (Fig. l). Hence elytral suture, a fungivore, fungus-eater; hence fungivorous (= mycophagous).
median ridge dividing the elytra symmetrically (Fig. 93, 94). fuscous, dark brown, approaching black.
empodium, a pad or extension between the tarsal claws. fusiform, spindle-shaped, i.e., broad at middle and narrowing
endemic, pertaining to species or higher taxa confined to a towards the ends.
geographical area under discussion; hence endemism. galea, outer lobe of maxilla (Fig. 5).

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gall, abnormal growth of plant tissue caused by various organ- hypogeal, subterranean (cf. epigeal).
isms, including insects. hypognathous (of the head), vertical, with mouthparts facing
gena (pl., genae), cheek, that part of head capsule on either downwards (Fig. 9, 10).
side below eye (Fig. 2, 4). hypomeron (pl., hypomera), epipleural fold, or in adult beetles
geniculate (of antennae), see p. 15 and Fig. 6, 28. (Polyphaga) lateral portion of pronotal disc extended ventrally
genital segment, modified 9th abdominal segment in beetles, on either side which meets lateral part of sternum (Fig. 3).
containing the genitalia; in the male the 10th segment is hypopharynx, a median lobe immediately behind mouth,
usually highly reduced or fused to segment 9. bearing taste receptors. Hence hypopharyngeal bracon (of
genitalia, structures relating to reproduction. beetle larvae), a transverse brace between hypopharynx and
gibbosity, irregular protuberance; hence gibbose, very anterior part of hypostomal margin.
crooked, twisted. hypopleurum (pl. hypopleura), in beetle Iarvae, area below
glabrous, without setae. ventrolateral suture.
globose, nearly spherical. hypostoma (pl., hypostomata), region of subgena behind
mandible. Hence hypostomal margin (of beetle larvae), a
glossa (pl., glossae), tongue; inner pair of lobes at apex of
ventral marginal thickening of the epicranial halves; hypo-
prementum of labium.
stomal tooth (in some groups of Curculionoidea), a tooth
Gondwana (`Land of the Gonds'), during much of the
situated in lower mouth between mandibles and base of
Mesozoic Era, a southern supercontinent consisting of maxillae.
continental blocks of South America, Africa, Madagascar,
incrassate (of antennae, etc.), thickened, suddenly swollen
India, Antarctica, and Australia with New Zealand.
towards the apex (Fig. 134).
grooming device, area of vestiture in a groove or impression
indigenous, originally inhabiting a particular geographical area
on inner face of protibia, usually for cleaning the antennae.
(but not necessarily restricted to that area).
gula, in prognathous insects, fused lower ends of post-occiput,
instar, life stage between moults in a nymph or larva,
forming a ventral plate (Fig. 2, 4). Hence gular suture, line of
numbered to designate developmental sequence; e.g., the first
division between gula and genae; ventro-anterior extension of
instar is the stage between the egg and the first moult.
postoccipital suture.
integument, outer layer of an insect, comprising epidermis and
haemolymph, watery body fluid in insects corresponding to
cuticle (see also exosketeton).
blood.
internal sac, part of the aedeagus.
habitus, general form and appearance.
intersegmentaI membrane, flexible, infolded membrane
hemisternites (of female genitalia), two sclerites of sternum 9
connecting segments of body and appendages which allows
surrounding vulva.
freedom of movement (Fig. 91).
heterogeneous, differing in kind, unlike in quality.
Jurassic, second period of Mesozoic Era, between Triassic and
heteromeroid (of trochanterofemoral junction), strongly Cretaceous, extending about 180-35 million years before the
oblique, so that there is direct contact between femur and coxa
.
present.
(Fig. 58, 59).
labium (pl., labia), lower lip, or fused second maxillae,
heteromerous, with the three pairs of tarsi having different forming floor of mouth in mandibulate insects, behind first
numbers of tarsomeres, e.g., tarsal formula 5-5-4. maxillae and opposed to labrum. Hence labial palp, a paired,
Hexapoda, superclass of phylum Arthropoda, including 1-4-segmented sensory appendage of labium.
Collembola, Protura, Diplura, and lnsecta, characterised by labrum, upper lip, joined lo clypeus in front of mouth (Fig. 1).
mouthparts consisting of mandibles, maxillae, and labium,
lacinia (pl. laciniae), bladelike inner lobe of maxilla, articu-
head with anterior and posterior tentorial arms, 3-segmented
lated to stipes and bearing brushes of hairs or spines (Fig. 5).
thorax, normally with 3 pairs of legs in the adult, and abdomen
with 6-I2 segments, with at most rudimentary limbs. lamella (pl. lamellae), a thin, platelike or leaflike process.
Hence lamellate (of antennae), with apical segments having
hinge mechanism, a structure of the medial loop of the hind
flat, leaflike projections on one side which sometimes cannot
wing used for wing folding (in Archostemata and Adephaga)
(Fig. 30) but usually can (Fig. 31) be brought closely together.
(Fig. 51).
lateroventral, lateral and below median horizontal plane.
Holarctic, faunal region comprising the northern part of the
Northern Hemisphere. ligula (pl., ligulae), collective name for glossae plus para-
glossae, whether fused or separate (Fίg. 2).
holometabolous, passing through a complete metamorphosis
in which the larva is very different from the adult and littoral, in freshwater ecosystems, in shallow water where light
transforms dramatically by means of a pupal stage. penetration extends to the bottom sediments, which are col-
onised by rooted plants; in marine ecosystems, intertidal,
humeral angle, anterolateral portion of elytra.
inhabiting the shore.
humerus (pl. humeri), shoulder, anterior basal portion of wing.
locking mechanism, a structure preventing movement, e.g.,
hydrophilous, water loving, living at least partly in water.
pronotal locking mechanism in adult ptilodactylid and
hyphae, threadlike strands making up the mycelium of a chelonariid beetles (Fig. 163, I64).
fungus.

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mala, a lobe of the mandible or jaw. l
nomina
nominal (of a species), bearing a name, but not necessarily a
Malaise trap, see p. 17. valid biological entity.
Malpighian tubes, internal excretory structures removing notopleural suture, in Adephaga, Archostemata, and
impurities from the body fluid. Myxophaga (Coleoptera), suture separating pronotal epipleuron
mandibles, first paired mouthparts, stout and jawlike in from pleuron (Fig. 2).
chewing insects (Fig. 1, 4). Hence mandibular socket, an notosternal suture, in Polyphaga, suture between hypomeron
impression or emargination containing mandibular base. of pronotum and prosternum where these plates meet directly
maxilla (pl., maxillae), second pair of jaws in insects with (i.e., not separated by an external pleuron).
chew-ing mouthparts; represented in all insects with functional notum (pl., nota), dorsal sclerite of a thoracic segment (Fig. 1 —
mouthparts (Fig. 5). Hence maxillary palps, paired, 1-5- pronotum).
segmented (usually 3 or 4 in Coleoptera) sensory appendages oblongum cell, feature of venation in hind wing of Adephaga
of the maxillae. (Fig. 46-48).
maxillolabial complex, a compound structure consisting of occiput, posterior part of epicranium between vertex and neck,
maxillary and labial elements (e.g., in larvae of Limnichidae). rarely present as a distinct sclerite (Fig. 6). Hence occipital
media, fifth longitudinal vein system (M), usually divided into carina, a well defined, elevated crest along hind margin of
a generally convex anterior branch (MA) and a concave occiput (e.g., in some Leiodidae).
posterior branch (MP) (Fig. 46, 47). ocellus (pl., ocelli), in adult insects, a simple eye consisting of
medial, near or at the middle of a structure or area. a single beadlike lens, occurring singly or in small groups; in
medial loop, see oblongum cell. holometabolous larvae, a stemma.
median lobe, part of aedeagus. ocular canthus, a lobe partially or completely dividing each
membranous, thin and more or less transparent (of wings); eye (e.g., in Gyrinidae, Tenebrionidae — Fig. 11, f).
thin and pliable (of cuticle). Oligocene, geological epoch, about 40-25 million years before
mentum (pl., menta), part of labium, proximal to prementum, the present.
attached to and sometimes fused with submentum (Fig. 2, 4). ommatidium (pl. ommatidia), a visual `cell' or element of the
meso-, prefix often pertaining to the mid le segment of the arthropod compound eye.
thorax, used as in `mesocoxa', `mesepimeron'. onisciform (of larvae), shaped like a slater (Oniscus sp.),
Mesozoic, geological era about 225-70 million years before depressed and broadly spindle-shaped; e.g., Elmidae.
the present. operculum (pl., opercula), a lid or cover; in larval Dryopidae a
meta-, prefix often pertaining to the hind segment of the ventral, flat cover to the chamber containing the retracted anal
thorax, as in `metacoxa', `metepimeron', gills.
opisthognathous (of head), with a posteroventral position of
metamorphosis, a radical change in form, especially the
mouthparts resulting from deflexion of the facial region.
change larva—pupa—adult in holometabolous insects.
orogeny, episode of mountain building in geological time.
microsculpture, rnicroscopic textural features of insect
integument. oviposit, to lay eggs.
microsetae, very small, short, hairlike structures; hence palpiger, a sclerite bearing a palp, esp. the palp-bearing
microsetose. structure of the mentum.
mola, ridged or roughened grinding surface of mandible, paraglossa (pl., paraglossae), a paired labial structure at apex
corres-ponding to subgalea of maxilla when the mandible is of prementum, connected at either side to glossae and either
compound. free or two-jointed; corresponding to galea of maxilla (Fig, 4,
moniliform (of antennae), see p. 14 and Fig. 19. 5).
monophagous, host-specific; restricted to one species of food paraphyletic (of a taxon or taxa), not including all the descen-
plant or animal (cf. polyphagous). dants of a common ancestor, i.e., not a natural' group.
monotypic (of a taxon), containing only one subordinate paraprocts, the halves of the ninth tergite.
taxon, e.g., a genus of only one species (cf. polytypic). parasitoid, an internal or external parasite that eventually kills
multisetose, bearing numerous setae. its host organism.
mycangium (pl., mycangia), any one of a variety of special paratergite, lateral marginal region of notum, lateral sclerite of
pocket-shaped receptacles used to carry symbiotic fungi, e.g., abdominal segments.
in bark beetles (Scolytinae). pectinate (of antennae), with one side of each segment
mycophagous, fungus-eating (= fungivorous). protruding greatly, forming a series of long, sharp points like
the teeth of a comb (Fig. 22).
myrmecophilous, ant-loving; applied to insects that live in ant
nests. pedicel, second segment of the insect antenna, supporting the
flagellum (Fig. 1, 6).
nacreous, having an iridescent lustre like mother of pearl.
peduncle, a stalk or stalklike structure, usually supporting an
niche, functional position of an organism in a community. organ or other structure; hence pedunculate.
nocturnal, active at night (cf. diurnal). phallobase, part of the aedeagus.

-94--
94-
pharyngeal process, a hollow membranous structure pterothorax, fused mesothorax and metathorax.
projecting into the oral cavity from the internobasal angle of pubescent, clothed with fine hairs.
the mandibles; present in nearly all Curculionoidea, absent or punctate, with puncture-Iike depressions, e.g., of the
only rudimentary in other beetles. integument.
phloem, vessels transporting dissolved nutrients from leaves to pupa, inactive transformative stage of holometabolous insects,
roots in vascular plants. intermediate between larva and adult; hence pupal cell, a
phreatic, underground, usually in reference to the moist cavity in the soil οr made of plant material or debris in which
habitat between the particles of coarse alluvial sediments. the larva pupates; puparium (pl. puparia), final-instar larval
phylogeny, evolutionary history of a group of organisms. integument within which pupation occurs.
phytophagous, feeding in/on plants; hence phytophage. pygidium (pl., pygidia), tergum of last visible abdominal
plasmodia, in myxomycete fungi, a motile mass of protoplasm segment; in many adult beetles, the segment left exposed by
characteristic of the growth phase. the termination of the elytra (Fig. 189, 238).
plesiomorphy, ancestral character state (cf. apomorphy). pygopod, paired footlike organ at hind end of abdomen of
pleurite, see pleuron. certain larvae, assisting locomotion.
pIeuron (pl., pleura), in adults of Adephaga, ventrolateral part pyriform, pear-shaped.
of thoracic wall meeting epipleuron Iaterally and sternum radial cell (Fig. 46), structure formed around a sclerotised,
medially; separated from pleuron by notopleural suture, and pigmented blood sinus (pterostigma), playing a part in folding
from sternum by pleurosternal suture (Fig. 2). the wing apex beneath the elytra.
plica (pl., plicae), a fold or wrinkle. radial loop, hind wing structure in beetles, consisting of the
plumose (of antennae), see p. 14. radial bar, which ends abruptly soon after entering the ptero-
stigma, and the pterostigma itself, transformed into the radial
posterolateral, towards the rear and to one side; similarly with
cell (Fig. 46-51).
-dorsal, -median, -ventral, etc.
postmedian, situated behind the middle. raptorial (of legs), adapted for seizing and holding prey (Fig.
1 10a, b).
postmentum, basal part of labium proximal to prementum.
recumbent, lying down, reclining.
postocular, behind the compound eyes.
relict, surviving in isolation.
predacious, preying upon other organisms, e.g., Carabidae,
most Staphylinidae. rhabdom, site of photoreception in the compound eye.
prementum, region of labium distal to mentum, formed by riparian, living on banks of watercourses.
palpigers and labial stipites. rostrum (pl., rostra), a snoutlike projection or rigid extension
prepygidium, part of last abdominal tergum separated from of the head, bearing the mouthparts at the end, e.g., in adult
pygidium by a transverse suture. weevils (Fig. 6, 7); hence rostrate.
pro-, prefix often pertaining to anterior segment of thorax, as scape, first or basal segment of insect antenna (Fig. 1, 6).
in procoxa, proepimeron. sclerite, any discrete plate of the body wall bounded by mem-
prognathous (of the head), horizontal, with mouthparts brane or by sutures.
directed forwards (Fig. 8). sclerolepidia, a row of pits with special setae or scales at the
metepisternal suture in some weevils.
pronotum, upper and dorsal part of prothorax (Fig. 1). Hence:
pronotal carina, e.g., basal transverse carina in Anthribidae sclerotisation, hardening of the cuticle, involving development
(Fig. 245, 246), and sometimes referring to lateral margin of of crosslinks between protein chains.
pronotum (Fig. 1); pronotal comb, a row of spines on posterior scrobes, grooves for reception or concealment of an
margin of pronotum, e.g., in Ptilodactylidae (Fig. 163); appendage; in adult Curculionoidea a pair of grooves, one on
pronotal epipleuron, in adult Adephaga, deflexed or inflexed either side of the rostrum, in which the antennae can rest (Fig.
portion of pronotum (Fig. 2). 7, d).
prosternum, sternum of the prothorax (Fig. 2, 3, 34-39). scutellum (pl., scutella), any small, shield-shaped plate;
Hence prosternal process, a posterior prolongation of posterior division of pterothoracic notum; in adult beetles,
prosternum behind procoxae. triangular piece at base of pronotum and between elytra (Fig.
prothorax, first thoracic segment, bearing anterior legs but no 1 ). Hence scutellary striole.
wings (Fig. 2, 3, 34-39). scutum (pl., scuta), second dorsal sclerite of mesothorax and
proximal, near or towards the anchored end of any appendage; metathorax, the middle division of the notum.
that part of a segment closest to the body (cf. distal). securiform, hatchet-shaped; e.g., the terminal segment of the
pseudotetramerous (of the tarsus), having apparently 4 con- palpi of some beetles (Fig. 15).
stituent articles, although 5 are actually present (Fig. 67-81). sensorium (pl., sensoria), a sensory receptor visible as a
pseudotrimerous (of the tarsus), having apparently 3 articles, circular opening covered by a membrane on antennae or legs.
although 4 are actually present (Fig. 80-82). sensory seta, a seta used as a simple sense organ, often a
pterostigma, pigmented spot on anterior margin of wing. contact or touch receptor.
seriate-punctate, with punctures arranged in striae.

-95-
 serrate (of antennae, etc.), see p. 14 and Fig. 20, 21. tegmen, part of the aedeagus.
sessile, attached directly, without any stem or petiole. tentorial pit, lateral depression in cranial wall usually near or
seta (pl. setae), a hairlike projection of the cuticle; hence at frontoclypeal suture (Fig. 1).
setiferous, bearing a seta or setae; setose, furnished with setae. tergite, see tergum.
sinuate (of edges and margins), wavy. tergum (pl., terga), upper or dorsal surface of any body
slime moulds, Mxomycetes, one of the four fungal phyla; segment of an insect, consisting of one or more sclerites.
hence slime flux, a plasma-like stage of slime moulds. terrestrial, living on land (cf. aquatic); sometimes taken as the
sooty moulds, fungi of the Ascomycetes: Capnodiaceae and complementary opposite of marine, i.e., including freshwater.
related families. testaceous, brownish yellow.
spiracle, external opening of a trachea. thorax, middle portion of insect body between head and abdo-
spore, among fungi, a microscopic structure functioning in men, consisting of three segments, each of which usually bears
dispersal and vegetative reproduction. a pair of articulated legs (Fig. 2).
spring mechanism, in Polyphaga, an elastic structure of the tibia (pl., tibiae), fourth segment of Ieg, between femur and
me dian loop used for folding the hind wing (Fig. 51). tarsus (Fig. 57). Hence tibial spur, a spinelike projection at or
spur, a thick cuticular appendage or spine connected to the near end of tibia.
body wall by a joint, especially on the tibia. tomentose, covered with short, matted, woolly hairs.
stemma (pl., stemmata), a simple eye of the often circular, tooth (in insects), a short, pointed process from an appendage
lateral eye-groups in holometabolous insect larvae. οr margin; a very stout spicule with a blunt apex.
sternite, see sternum. trachea (pl., tracheae), a spirally ringed, internal elastic air
sternum (pl., sterna), entire ventral division of any body tube in insects; an element of the respiratory system.
segment. trilobed (of the aedeagus), consisting of a ventrally sclerotised
stipes (pl., stipites), basic sclerite of maxilla, articulating partly phallobase to which a pair of freely articulated parameres are
with head, partly with cardo, and bearing galea, lacinia, and attached, and a simple penis lying above it; generally
maxillary palp. considered to be the most primitive configuration in beetles.
stria (pl., striae), in general, any fine longitudinal impressed trochanter, second segment of insect leg, between coxa and
line; in beetles, a longitudinal depressed line or furrow, often femur (Fig. 57-66).
punctured, usually extending from the base to the apex of the trochanterofemoral, pertaining to trochanter and femur con-
elytra; hence striate. jointly.
stridulatory apparatus, structure used in making sounds, trochantin, a small sclerite separated from pleuron by a mem-
consisting of a file-like organ and an opposing scraper or rasp. brane and connected by articular condyle with coxa (Fig. 57).
striole, rudimentary or reduced stria. trophic, pertaining to food or eating.
stroma (pl. stromata), mass of vegetative fungal hyphae truncate, cut off squarely at the tip.
bearing fruit bodies and/or spores. tubercle, a small, knoblike or rounded protuberance; hence
stylus (pl. styli), a small, pointed, non-articulated process. tuberculate.
sub-, prefix denoting under, slightly less than, or not quite. Tulgren funnel, funnel-shaped device for sorting Iiving insects
subcortical, beneath the bark of a tree. from organic debris by means of light and heat.
submentum, basal part of labium by which it is attached to urogomphi (sing., urogomphus), in larval beetles usually
head capsule (Fig. 2). paired posterior processes on tergum of ninth abdominal
segment, either jointed and movable by muscles or unjointed
sulcus, a groove or suture marking the line of fusion of two
and immovable.
formerly distinct plates or structures.
UV light-trap, see p. 17.
suture, a seam or impressed line indicating where two sclerites
join. Hence sutural flanges, posterior sutural edges of elytra, venter, entire undersurface of body of an animal; hence
separated by the sutural gap (Fig. 93). ventral.
synapomorphy, a shared derived character. ventrad, towards the venter.
systematics, study of biological classification based on ventrite, sternal sclerite of abdomen; the first ventrite is equi-
relationship through inherent similarity; hence systematic valent to the first visible basal sternite.
sequence of taxa, as used in this volume. vertex, top of head between eyes, frons, and occiput, anterior
tactile, pertaining to sense of touch, or used for touching. to occipital suture.
tarsus (pl., tarsi), terminal leg segment attached to apex of vesicle, a small sac or bladder.
tibia, consisting of 1-5 articles or tarsomeres. Hence tarsal waist, basal constriction of pronotum in beetles, e.g.,
formula, the number of tarsomeres on the protarsus, Anthicidae (Fig. 228-231).
mesotarsus, and metatarsus. weevil, beetle of the family Curculionidae.
taxonomy, theoretical study of classification, including its window trap, see p. 17.
bases, principles, procedures, and rules. wireworm, soil-inhabiting larva of Elateridae feeding on roots.

-96-
 KEYS TO FAMILIES OF COLEOPTERA Suborder Polyphaga
KNOWN FROM NEW ZEALAND 5(1) Antennae with a lamellate, flabellate, or exceptionally
pectinate club (Fig. 30-33) ... (p. 98) .. Group A
—Antennae not as above .. 6
I: PYRAMIDAL KEY
[provided by J. Klimaszewski] 6(5) Tarsi 5-5-5-segmented but appearing 4-4-4-seg-
1 First ventrite divided by metacoxae (Fig. 88, 89); mented, with segment 4 reduced and segment 3
prothorax usually with distinct notopleural sutures, distinctly bilobed, not so in some Chrysomelidae
distinct from sharp lateral margins (Fig. 2); tarsi 5- (exception: weevil groups Belidae : Aglycyderinae
segmented; antennae usually filiform (Fig. 17, 18); and Brentidae : Apioninae (Neocyba), in which tarsi
medial loop of hind wing with a hinge mechanism appear 3-3-3-segmented but then head with a rostrum)
(Fig. 51) ... (p. 20) ADEPHAGA .. 2 (Fig. 67-81) ... (p. 98) .. Group B
—First ventrite usually undivided by metacoxae and —Tarsi not as above, if 5-5-5-segmented with segment 4
continuous from side to side (Fig. 90); prothorax reduced then all segments simple, and if appearing 3-
without notopleural sutures — ventral portion of notum 3-3-segmented then head without a rostrum ... 7
(hypomeron) jointed direcly to sternum on either side
7(6) Body bullet-shaped, with sides subparallel and
by notosternal suture (Fig. 3); tarsi and antennae often
not as above; medial loop of hind wing with a spring abdominal apex triangular; head deflexed; abdomen
with first 2 ventrites strongly connate, the suture
mechanism (Fig. 51) ... (p. 23) POLYPHAGA .. 5
partially obliterated; metasternum with a distinct
transverse suture ... (Fig. 157) .. [19] BUPRESTIDAE
—Body not so; first 2 ventrites rarely fused ... 8
Suborder Adephaga
Superfamily Caraboidea 8(7) Tarsi usually 5-5-5-segmented, with terminal seg-
2(1) Terrestrial beetles with legs adapted for crawling or ment enlarged and ending in strong claws, often longer
running; metacoxae not meeting elytra laterally (Fig. than all other segments combined (Fig. 160); aquatic
2); antennae largely pubescent; dorsal surface with or semi-aquatic beetles broadly oval in outline,
pronounced, erect sensory setae (Fig. 107) ... 3 convex, with deflexed head more or less concealed by
—Aquatic beetles with at least hind legs adapted for pronotum from above; legs retractable into body
swimming, often flattened and with dense pubescence cavities (Fig. 52, 53); prosternum with a process
(Fig. 108-110); metacoxae extending laterally to meet fitting into a cavity of mesosternum (Fig. 52, 53);
elytra (Fig. 88, 110, b); antennae entirely glabrous, or antennae weakly clubbed (3-7 segments), comblike,
nearly so; dorsal surface without pronounced, erect sawtoothed, or distinctly clubbed (Fig. 29)
sensory setae (phreatic species, e.g., Kuschelydrus, ... (p. 99) .. Group C
with long, fine, setae on sides — Fig. 109) ... 4 —Tarsi other than 5-5-5, or if 5-5-5-segmented then
terminal segment not enlarged; sometimes segment 4
reduced and tarsi appearing 4-4-4-segmented (Phalac-
3(2) Metacoxae widely separated; antennae thick, mon-
ridae); other features not as in above combination... 9
iliform (Fig. 19); pronotum usually with 3 deep
longitudinal grooves, or at least 1 groove; head with a 9(8) Maxillary palps usually longer than antennae (Fig.
distinct neck; metasternum without transverse suture 112, 116) (exception: Meropathus, with maxillary
... (Fig. 101) .. [1] RHYSODIDAE palps much shorter than antennae — Fig. 117); an-
—Metacoxae contiguous or narrowly separated (Fig. 2); tennae 7-11-segmented with a 2-5-segmented club;
antennae slender, filiform (Fig. 1); pronotum without hydrophilous species, spherical or moderately convex
deep grooves; head usually without a distinct narrow to ovoid and flattened ... (p. 99) .. Group D
neck; metasternum with a transverse suture (Fig. 2) —Maxillary palps shorter than antennae, inconspicuous;
... (Fig. 102-107) .. [2] CARABIDAE antennae usually 11-segmented, with a 3-segmented
club or unclubbed; shape usually not as above ... 10
4(2) Antennae longer, filiform; eyes not divided into two;
forelegs short; hind legs long, modified for swimming, 10(9) Pronotum with posterolateral angles strongly pro-
flattened and fringed with hairs duced behind (Fig. 165-167); body elongate and
... (Fig. 108, 109) .. [3] DYTISCIDAE narrowly subparallel (Fig.165-167); click mechanism
—Antennae short, with a large pedicel; eyes each divided present, consisting of a prosternal process (Fig. 54, a)
into two; forelegs long, raptorial; middle and hind legs which fits into a deep mesosternal cavity (Fig. 54, b);
short and modified for swimming antennae filiform, serrate, pectinate, flabellate, or in-
... (Fig. 110) .. [4] GYRINIDAE crassate ... (p. 99) .. Group E

—97—
 —Pronotum and body shape not sas above; click mech- -Tarsal formula 2-2-2, 3-3-3, 3-4-4, 4-4-4, or 5-5-5, if
anism absent; antennae often clubbed ... 11 4-4-4 then canthus absent and first 3 ventrites not
fused together ... 17
11(10) Elytra short, truncate, exposing at least 2 ab-
dominal tergites, usually more, the exposed tergites 17(16) Tarsal formula 5-5-5 (excluding those families
well sclerotised rather than membranous; abdomen with 5-5-5-segmented tarsi already keyed out)
usually flexible ... (p. 99) .. Group F ... (p. 102) .. Group J
—Elytra not so, broadly rounded posteriorly and usually —Tarsal formula 2-2-2, 3-3-3, 3 4 4, or 4-4-4; antennae
covering entire abdomen, but if shorter then exposing filiform, moniliform, or with a distinct, compact, l-3-
only the abdominal apex; tergites membranous or segmented club; mostly small beetles of average
weakly sclerotised ... 12 length 2.5-3 mm, but some up to 17 mm long
... (p. 104) .. Group K
12(11) Antennae with an abrupt 3- or 4-segmented club,
geniculate, usually retractable into a cavity on under-
side of prothorax; body oval, disc-shaped, moderately Group A
to strongly convex, glossy, usually glabrous and black; Includes families with a lamellate or flabellate (exception-
elytra shortened, exposing the pygidium; length 2-9 ally pectinate) antennal club.
mm ... (Fig. 114, 115) .. [6] HISTERIDAE
—Antennae and body not as above ... 13 SCARABAEOIDEA: Lucanidae, Trogidae, Scarabaeidae.
BOSTRICHOIDEA: Bostrichidae (Euderia). BYRRHOIDEA:
13(12) Body broadly oval, strongly convex, glossy; elytra Chelonariidae (Βrοunia). TENEBRIONOIDEA: Rhipiphoridae.
usually with 2 fine grooves near suture; in lateral view
basal part of elytra and pronotum angular; antennae Al Lamellae of antennal club not capable of close
with a 3-segmented club; tarsi 5-5-5- or 5-5-4-seg- apposition (Fig. 30) ... 2
mented but appearing 4-4-4-segmented because of —Lamellae of antennal club capable of close apposition
vestigial segment 4 ... (Fig. 196) .. [48] PHALACRIDAE (Fig. 31) .3
—Body not as above; elytra usually without 2 grooves
near suture; in lateral view basal part of elytra and Α2(1) Antennae geniculate (exception: Ceratognathus
pronotum not distinctly angular; antennae and tarsi not parrianus) ... (Fig. 139-143) .. [13] LUCANIDAE
as above ... 14 —Antennae not geniculate ... 4

14(13) Frons with 1 or 2 ocelli ... (p. 100) .. Group G Α3(1) Elytra dull, more or less tuberculate, without striae
—Frons without ocelli ... 15 ... (Fig. 144) .. [14] TROGIDAE
— Elytra usually glossy, glabrous, or semiglabrous, or
15(14) Body loosely articulated, elongate, oblong to sub- with velvety pubescence (e.g., Costelytra), and often
parallel; integument soft, often velvety in appearance with striae ... (Fig. 145-152) .. [15] SCARABAEIDAE
(e.g., Fig. 169, 186, 223), with decumbent pubescence
and often additional scattered and erect setae, or with Α4(2) Metatarsi 4-segmented
short and inconspicuous pubescence; pronotum and ... (Fig. 214) .. [64] RHIPIPHORIDAE
elytra sometimes with ridges divided into cell-shaped - Metatarsi 5-segmented ... 5
compartments (Fig. 168) ... (p. 100) .. Group H
—Body compact; integument hard, glabrous or with Α5(4) Antennal club with more than 3 lamellae (up to 7)
decumbent pubescence; pronotum and elytra not as ... (Fig. 164) .. [26] CHELONARIIDAE (Brounia)
above ... 16 —Antennal club of male with 3 flabella
... (Fig. 174) .. [35] BOSTRICHIDAE (Euderia)
16(15) Tarsal formula 5-5-4, exceptionally 4-4-4 in some
Tenebrionidae, but then base of each antenna ob-
scured from above by a shelf-like expansion (canthus Group Β [assisted by G. Kuschel]
— Fig. 11, f) and first 3 ventrites fused together; body Includes families with tarsi 5-5-5-segmented but appear-
broadly to narrowly oval, medium-sized to large (l.5- ing 4-4-4-segmented because of vestigial segment 4, and
20 mm); procoxae usually conical and projecting; segment 3 distinctly bilobed. Exception: Aglycyderinae
antennae 11-segmented (10-segmented in Archaeo- (family Belidae) and Neocyba (Brentidae: Apioninae), in
glenes, Tenebrionidae), filiform, serrate, incrassate, which tarsi appear 3-3-3-segmented but are 4-4-4-seg-
sawtoothed, comblike, or flabellate, without a club or mented, and head with a rostrum.
with a loosely defined, elongate, 3- or 4-segmented
club ...(p. 101) .. Group I CHRYSOMELOIDEA: Chrysomelidae (incl. Bruchinae), Ceram-

— 98—
bycidae. CURCULIONOIDEA: Nemonychidae, Anthribidae, ments combined, and ending in strong claws (exception:
Belidae (incl. Aglycyderinae), Brentidae (incl. Apioninae), some Limnichidae (Hyphalus) and some Byrrhidae, with
Curculionidae (including Brachycerinae, Curculioninae, 4-4-4-segmented tarsi); body broadly oval and convex,
Rhynchophorinae, Cossoninae, Scolytinae, Platypodinae). deflexed head concealed by pronotum from above, legs
retractable into body cavities, prosternum with a process
B1 Anterior part of head never with a rostrum 2 fitting into a cavity of mesosternum, and antennae with a
—Anterior part of head elongate, forming a long or short weak club of 3-7 segments; aquatic or semi-aquatic.
rostrum (exceptionally short in Scolytinae, Platy-
podinae), with modified mouthparts (Fig. 6, 7) ... 3 BYRRHOIDEA: Byrrhidae, Dryopidae, Elmidae, Limnichidae.

Β2(1) Antennae usually inserted on frontal prominences Cl Antennae shorter than pronotum, comblike, often
and capable of being flexed backwards over body, at concealed ... (Fig. 159) .. [21] DRYOPIDAE
least two-thirds as long as body —Antennae usually longer than pronotum, slender, saw-
... (Fig. 234-237) .. [76] CERAMBYCIDAE toothed, sometimes weakly clubbed ... 2
—Antennae not inserted on frontal prominences, not
capable of being flexed backwards, usually only mod- C2(1) Body narrowly elongate; elytra with lateral mar-
erately elongate (exception: Cryptocephalinae, with gins subparallel ... (Fig. 160) .. [22] ELMIDAE
long antennae capable of being flexed backwards) —Body broadly oval; elytra with lateral margins broadly
... (Fig. 238-243) .. [77] CHRYSOMELIDAE curved (Fig. 158, 161) .. 3

B3(l) Antennae geniculate, with a more or less compact C3(2) Hind margin of pronotum evenly curved or slightly
club (Fig. 6, 28); scape long sinuate; frontoclypeal suture absent; body glabrous or
... (Fig. 251-260) .. [82] CURCULIONIDAE semiglabrous ... (Fig. 158) .. [20] BYRRHIDAE
—Antennae straight, with or without a club (Fig. 244- —Hind margin of pronotum strongly sinuate; fronto-
clypeal suture present; body densely pubescent
249) ... 4
... (Fig. 161) .. [23] LIMNICHIDAE
Β4(3) Labrum separated from clypeus by a distinct suture
(Fig. 6, b); mandibles with a mola ... 5 Group D
—Labrum fused with clypeus, without a suture; mandi-
Includes families with hydrophilous species commonly
bles without a mola ... 6
bearing extremely elongate maxillary palps, usually longer
than the antennae, and antennae 7-11-segmented with a
Β5(4) Pronotum without carinae as below; antennal seg-
2-5-segmented club.
ment 3, 4, or 5 extending to eye; spurs present on all
HYDROPHILOIDEA: Hydrophilidae. STAPHYLINOIDEA: Hyd-
tibiae; all 5 ventrites free
raenidae.
... (Fig. 244) .. [78] NEMONYCHIDAE
—Pronotum usually with a transverse basal carina and a
BDodyusal1phericbodyvalnstrg
lateral carina; antennal segment 1 or 2 extending to
convex; maxillary palps usually shorter than antennae;
eyes; spurs absent on all tibiae; only 5th ventrite
antennae with a 3-segmented pubescent club
completely free, other ventrites partly fused or firmly
... (Fig. 111-113) .. [5] HYDROPHILIDAE
braced ... (Fig. 245, 246) .. [79] ANTHRIBIDAE
—Body usually moderately narrowly elongate, rarely
more broadly rounded; maxillary palps often longer
Β6(4) Rostrum moderately elongate and narrow; protibia
than antennae (exception: Meropathus species, with
with a grooming device on inner face opposite tarsal
maxillary palps much shorter than antennae); antennae
articulation, consisting of fine and dense pubescence
usually with a 5-segmented pubescent club
in a broad, shallow groove or impression; gular sut-
... (Fig. 116, 117) .. [7] HYDRAENIDAE
ures distinctly separate or obsolete
... (Fig. 247, 248) .. [80] BELIDAE
—Rostrum usually extremely elongate (Fig. 249); pro- Group Ε
tibia without a grooming device; gular sutures merged
Includes families with posterolateral angles of pronotum
into one and always distinct
strongly produced posteriorly, body narrowly elongate
... (Fig. 249, 250) .. [81] BRENTIDAE
and subparallel, clicking apparatus present, and antennae
filiform, serrate, pectinate, flabellate, or incrassate.
Group C
ELATEROIDEA: Elateridae, Eucnemidae.
Includes families with 5-5-5-segmented tarsi, each ter-
minal segment enlarged, often longer than all other seg- Ε1 Labrum concealed beneath clypeus, externally not

-99 —
 visible; ventrites 4 and 5 connate F7(6) Antennae filiform, serrate, or pectinate but never
... (Fig. 165) .. [27] EUCNEMIDAE with a distinct club ... (Fig. 169) .. [30] CANTHARIDAE
—Labrum visible; ventrites 4 and 5 not connate (in part; see also Group H)
... (Fig. 166, 167) .. [28] ELATERIDAE —Antennae with a distinct, compact club (Fig. 23)
... (Fig. 187) .. [42] NITIDULIDAE (in part)

Group F
Includes families with short and usually truncate elytra, Group G
exposing at least 2 but usually more abdominal tergites, Includes families with frons bearing 1 or 2 ocelli.
the exposed tergites well sclerotised, and abdomen usually
flexible. Family Ptiliidae, with some species having trun- BOSTRICHOIDEA: Dermestidae (in part). DERODONTOIDEA:
cate elytra and 2 abdominal tergites exposed, is included in Derodontidae. STAPHYLINOIDEA: Staphylinidae (Microsil-
Group K (p. 104). phinae, Omaliinae, some Proteininae).

CHRYSOMELOIDEA: Chrysomelidae (Bruchinae). CUCUJOIDEA: G1 Head with 1 ocellus on frons, positioned medially
Nitidulidae (in part; see also Group J), Monotomidae (in ... (Fig. 173) .. [34] DERMESTIDAE (in part)
part; see also Groups J, K). ELATEROIDEA: Cantharidae (in ... [12] STAPHYLINIDAE (some Proteininae)
part; see also Group H). STAPHYLINOIDEA: Staphylinidae —Head with 2 ocelli on frons, positioned laterally (Fig.
(except Microsilphinae and some Proteininae of genus 123, 124) ... 2
Silphotelus). TENEBRIONOIDEA: Salpingidae (Inopeplinae
only), Rhipiphoridae (see also Group A). G2(l) Head and pronotum not strongly sculptured; body
smooth and glabrous or with sparse pubescence and
F1 Elytra usually truncate posteriorly, exposing at least 3 punctation ... (Fig. 123, 124) .. [12] STAPHYLINIDAE
but most often 6 or 7 abdominal tergites ... 2 (Microsilphinae, Omaliinae, some Proteininae)
—Elytra truncate or rounded posteriorly, exposing 2 or —Head with a series of canals and bridges; pronotum
rarely 3 abdominal tergites ... 4 divided by longitudinal ridges into several cell-like
depressions ... (Fig. 170) .. [31] DERODONTIDAE
F2(1) Abdomen short, broad, inflexible; head, pronotum,
and often elytra with setose foveae
.. (Fig. 126) .. [12] STAPHYLINIDAE (Pselaphinae) Group Η
—Abdomen more elongate, narrow to moderately broad, Includes families with soft-bodied species having decum-
flexible; usually 6 or 7 tergites exposed ... 3 bent pubescence and often additional scattered and erect
setae, or with inconspicuous pubescence, and body sur-
F3(2) Metatarsi 4-segmented; tarsal formula 5-5-4 face often with a velvety appearance.
... (Fig. 227) .. [72] SALPINGIDAE (Inopeplinae)
— Metatarsi not 4-segmented, or if so then tarsal formula BYRRHOIDEA: Ptilodactylidae. CLEROIDEA: Cleridae, Mely-
other than 5-5-4 ridae. ELATEROIDEA: Cantharidae (in part; see also Group
... (e.g., Fig. 125) .. [12] STAPHYLINIDAE (in part) F), Lycidae. EUCINETOIDEA: Scirtidae (=Helodidae).
TENEBRIONOIDEA: Oedemeridae, Pyrochroidae, Scraptiidae.
F4(1) Antennae appearing 10-segmented, with usually a
1-segmented club; elytra parallel-sided, usually with H1 Tarsal formula 5-5-5 ... 2
rows of punctures, truncate posteriorly, leaving pygid- —Tarsal formula 5-5-4 ... 7
ium exposed ... (Fig. 188, 189) .. [43] ΜΟΝΤΙDAΕ
(Lenax, Monotoma) H2(1) Body flattened, loosely articulated; pronotum with
—Antennae usually 11-segmented and not as above; several ridges dividing disc into irregularly shaped
elytral shape not as above ... 5 cell-like depressions; elytra leathery, with strong,
depressed punctures between longitudinal ridges
F5(4) Tarsi appearing 4-4-4-segmented, with segment 3 ... (Fig. 168) .. [29] LYCIDAE
bilobed; body robust, egg-shaped, usually broadest —Body form not as above; pronotum without ridges and
towards apex; head often concealed from above irregularly shaped cell-like depressions; elytra hard,
... (Fig. 238) .. [77] CHRYSOMELIDAE (Bruchinae) with or without longitudinal ridges, but never with
—Tarsi 5-5-5-segmented; body shape not as above ... 6 strongly impressed punctures between them ... 3

F6(5) Antennae flabellate; body wedge-shaped Η3(2) Scutellum heart-shaped; hind margin of pronotum
... (Fig. 214) .. [64] RHIPIPHoRIDAE (see also Group A) with minute, blunt teeth forming a transverse comb
—Antennae and body shape not as above ... 7 ... (Fig. 163) .. [25] PTILODACTYLIDAE

— 100 —
 —Scutellum subtriangular; hind margin of pronotum I1 Body moderately to strongly flattened (Fig. 192-195),
without such teeth ... 4 often hairless; antennae filiform or beadlike, some-
times with a loose 2-4-segmented club; mandibles
Η4(3) Penultimate tarsal segment simple, never bilobed more or less pointing forwards; elytra usually striate
... (Fig. 186) .. [41] MELYRIDAE (=Dasytidae) 2
... (Fig. 183) .. [39] CLERIDAE (Metaxina) —Body and antennae not as above; mandibles usually
—Penultimate tarsal segment bilobed ... 5 not pointing forwards; elytra striate or not ... 4

Η5(4) Body often subcylindrical, usually covered with 12(1) Body moderately flattened, appearing pubescent;
bristly hairs; head broad, as broad as pronotum or tarsal segment 2 and/or 3 lobed below or, if not lobed,
broader; pronotum much narrower than elytra; ant- then antennal scape at least as long as 3 following seg-
ennae often with a loose 3-segmented club ments combined; pronotum without sublateral ridges,
... (Fig. 184) .. [39] CLERIDAE often with anterior angles produced; antenna with or
—Body flattened to moderately convex, lacking bristly without club (Fig. 192, 193) [45] SILVANIDAE
hairs (pubescence short and decumbent); head usually —Body strongly flattened, usually glabrous (Fig. 194,
narrower than pronotum; pronotum as broad as elytra 195); tarsal segments 2 and 3 not lobed below; anten-
or slightly narrower; antennae without a 3-segmented nal scape as long as or shorter than the 2 following
club ... 6 segments combined; pronotum usually with sublateral
carinae; antennae without a club ... 3
Η6(5) Labrum sclerotised and distinct, visible from
above; head with characteristic keels below eyes (Fig. 13(2) Pronotum without 2 sublateral carinae
12) ... (Fig. 153, 154) .. [16] SCIRTIDAE ... (Fig. 194) .. [46] CUCUJIDAE
—Labrum membranous and covered by clypeus, not —Pronotum with 2 sublateral carinae
visible from above ... (Fig. 169) .. [30] CANTHARIDAE ... (Fig. 195) .. [47] LAEMOPHLOEIDAE

Η7(1) Eyes not projecting, C-shaped, strongly emarginate; 14(1) Base of each antenna covered by a canthus (Fig. 11,
pronotum usually with 2 small basal impressions f); abdominal sternites 1-3 fused together
... (Fig. 233) .. [75] SCRAPTIIDAE ... (Fig. 219-221) .. [68] TENEBRIONIDAE
—Eyes prominent, slightly to strongly projecting, slight- —Base of antennae exposed; abdominal sternites 1-3 not
ly emarginate or not; pronotum usually without 2 basal fused together ... 5
impressions ... 8
15(4) Pronotum distinctly narrower at base than elytra,
Η8(7) Eyes not strongly projecting laterally; only pen- half to two-thirds of elytral width (Fig. 228-232)... 6
ultimate tarsal segment bilobed; antennae usually fili- —Pronotum usually as wide at base as elytra, or nearly so
form ... (Fig. 223) .. [70] OEDEMERIDAE (exception: Ulodidae, with pronotum much narrower
—Eyes strongly projecting laterally; penultimate and basally than elytra– Fig. 217) ... 9
antepenultimate tarsal segments lobed; antennae usu-
ally pectinate or serrate 16(5) Pronotum without lateral margins; body ant-shaped
... (Fig. 224, 225) .. [71] PYROCHROIDAE (Fig. 228-232) ... 7
—Pronotum with lateral margins; body not as above ... 8

Group I I7(6) Ventrites all free; penultimate tarsal segment lobed;

eyes with fine facets, not notched near antennal base;
Includes families with 5-5-4-segmented tarsi, exceptionally
4-4-4-segmented in some Tenebrionidae (but then base of elytral epipleura incomplete
antenna covered by a canthus), procoxae usually conical ... (Fig. 228-231) .. [73] ANTHICIDAE
—Basal 2 ventrites connate; penultimate tarsal segment
and projecting, and antennae without or with a loosely
defined, elongate, 3- or 4-segmented club. small, simple, and antepenultimate segment lobed
beneath; eyes coarsely faceted, slightly notched near
CUCUJOIDEA: Cucujidae, Laemophloeidae, Silvanidae (in antennal base; elytral epipleura absent
part; see also Group J). TENEBRIONOIDEA: Aderidae, ... (Fig. 232) .. [74] ADERIDAE (= Euglenidae)
Anthicidae (incl. Pedilinae), Archeocrypticidae, Chalco-
dryidae, Melandryidae, Mordellidae, Phloeostichidae 18(6) Pronotum and elytra with fine and dense punc-
(males only; females in Group F), Salpingidae (excl. tation; antennae slightly incrassate, but without a
Inopeplinae: see Group F), Tenebrionidae, Ulodidae distinct club; body form not as below
(formerly as Zopheridae). ... (Fig. 190) .. [44] PHLOEOSTICHIDAE
(males of Agapytho; see also Group J)

-101-
 —Pronotum usually coarsely punctate, and elytra with Chaetosomatidae, Phycosecidae, Trogossitidae (incl.
rows of coarse punctures; antennae with a 3-seg- Protopeltinae). CUCUJOIDEA: Cavognathidae, Cryptophag-
mented, loosely formed club; often resembling small idae (in part; males with tarsi 5-5-4 are in Group I),
carabid beetles ... (Fig. 226) .. [72] SALPINGIDAE Cucujidae, Laemophloeidae, Silvanidae (in part; see also
(excl. Inopeplinae) Group I), Erotylidae, Languridae, Nitidulidae (in part; see
also Group F), Phloeostichidae, Monotomidae (in part;
I9(5) Antennae with a 3-segmented, elongate club ... 10 see also Groups F and K). EUCINETOIDEA: Eucinetidae.
—Antennae filiform, serrate, or slightly incrassate, with- STAPHYLINOIDEA: Agyrtidae, Leiodidae, Scydmaenidae.
out a distinct 3-segmented club ... 11
J1 Pronotum hood-like in lateral view, partially covering
I10(9) Body irregularly oval in outline; pronotum and head from above; antennae, if clubbed, then with all
elytra with margins not as below segments long and club not symmetrical; body shape
... (Fig. 191) .. [44] PHLOEOSTICHIDAE from elongate cylindrical to oval
(males of Priasilpha) ... (Fig. 177) .. [36] ANOBIIDAE (excluding Ptininae)
—Body approximately oval in outline; pronotum and —Pronotum not as above; antennae, if clubbed, then with
elytra with distinct sharp margins club symmetrical (exception: some Dermestidae);
... (Fig. 210) .. [60] ARCHEOCRYPTICIDAE body shape rarely cylindrical ... 2

I11(9) Body broadly oval, flattened, with decumbent or J2(1) Antennae not clubbed ... 3
erect pubescence, bristles, or scales forming patterns; —Antennae with a more or less distinct, 1-8-segmented,
pronotum narrower than elytral base, often with api- elongate and loosely formed or compact club; if club
cal angles produced ... (Fig. 217) .. [66] ULODIDAE not distinct then segment 8 smaller than adjoining
(formerly as Zopheridae) segments ... 8
—Body narrowly elongate, convex to slightly flattened,
with decumbent pubescence often forming patterns; J3(2) Body appearing spider-like; legs and antennae long
pronotum as wide as elytral base, with apical angles and thin; antennal insertions close together; body
rounded but not produced ... 12 covered with dense pubescence or scales forming
patterns ... (Fig. 178) .. [36] ΑΝΟΒΙΙDΑΕ
I12()Bodynarwlsubpe,ftnd;hao (Ptininae)
deflexed, not deeply inserted into prothorax; tibiae —Body elongate or oval; legs and antennae moderately
without combs or spurs long; antennal insertions distant from each other ... 4
... (Fig. 218) .. [67] CHALCODRYIDAE
—Body wedge-shaped, tapering posteriorly, convex; J4(3) Body elongate-oval, narrowed posteriorly; head
head deflexed, deeply inserted into prothorax; tibiae concealed from above, resting against procoxae in nat-
often with serrate combs or spines (Fig. 84) ... 13 ural position; metacoxae greatly expanded, covering
much of 1st ventrite; elytra often with cross-striations
I13(12) Last sclerotised tergite usually covered by elytra ... (Fig. 155) .. [ 17] EUCINTDAE
and not forming an acute process extending beyond —Body broadly to narrowly elongate, subparallel, often
elytra; head not abruptly constricted behind eyes; flattened; head usually prognathous, exposed and not
pronotum with lateral carinae incomplete; metacoxae concealed by pronotum; metacoxae not enlarged;
not plate-shaped ... (Fig. 212) .. [62] MELANDRYIDAE elytra without cross-striation ... 5
—Last sclerotised tergite produced posteriorly and
forming an acute process extending beyond elytra; J5(4) Body with long and erect hairs, sparsely and
head abruptly constricted behind eyes; pronotum with coarsely punctate, moderately flattened; elytra not
lateral carinae complete; metacoxae enlarged, plate- striate ... (Fig. 182) .. [38] CHAETOSOMATIDAE
shaped ... (Fig. 213) .. [63] MORDELLIDAE —Body without long and erect hairs, glabrous or with
short, sparse pubescence and coarse punctation; elytra
usually striate ... 6
Group J
Includes families with tarsal formula clearly 5-5-5 not J6(7) Body moderately flattened, usually pubescent;
already listed (see Groups B, C, D, F, and H). tarsal segment 2 and/or 3 lobed below or, if not lobed,
then antennal scape at least as long as the 3 following
BOSTRICHOIDEA: Anobiidae, Bostrichidae (in part, incl. segments combined; pronotum without sublateral car-
Lyctinae; see also Group A), Dermestidae (in part; see also inae ... (Fig. 192, 193) .. [45] SILVANIDAE
Group G), Jacobsoniidae (in part; see Group K for species —Body strongly flattened, appearing glabrous; tarsal
with tarsi 3-3-3), Nosodendridae, Ptinidae. CLEROIDEA: segments 2 and 3 not lobed below; antennal scape as

-102-
 long as or shorter than the 2 following segments J14(13)
J14(13) Body surface glabrous or with inconspicuous
combined ... 7 pubescence, usually strongly glossy ... 15
—Body surface moderately to densely pubescent, matt or
J7(6) Pronotum without 2 sublateral carinae slightly glossy ... 18
... (Fig. 194) .. [46] CUCUJIDAE
—Pronotum with 2 sublateral carinae J15(14) Body narrowly elongate, subparallel to cylindri-
... (Fig. 195) ... [47] LAEMOPHLOEIDAE cal; pronotum slightly to distinctly elongate ... 16
—Body broadly to narrowly oval; pronotum transverse
J8(2) Antennae with a 3-5- or 7-segmented club and and narrow, as wide as elytral base ... 17
usually with segment 8 smaller than adjoining ones
(Fig. 24), or if not so then body densely pubescent; J16(15) Body narrowly elongate, not cylindrical, sub-
often glabrous beetles; legs usually bearing strong parallel; pronotum without rasplike teeth at front;
spines ... (Fig. 120, 121) .. [10] LEIODIDAE metasternum extremely elongate, as long as all 5 ven-
—Antennae usually with segment 8 normally developed; trites combined or longer; body length 0.8-2 mm
usually pubescent beetles; legs without spines or with ... (Fig. 171) .. [32] JACOBSONIIDAE
small spines ... 9 —Body cylindrical or narrowly elongate and slightly
flattened; pronotum often with rasplike teeth at front;
J9(8) Body oblong to broadly oval, robust, with short, antennal club often asymmetrical (enlarged to one
dense pubescence or with erect hairs or scales often side); metasternum not extremely elongate, much
forming patterns; lower surface often with close, silky shorter than all 5 ventrites combined; body length 2.8-
pubescence; elytra if striate then with a scutellary 6 mm .... (Fig. 174-176) .. [35] BOSTRICHIDAE
striole; antennae with club 3-8-segmented, usually 3- (incl. Lyctinae; see also Group A)
segmented ... (Fig. 173) .. [34] DERMESTIDAE
—Body shape variable, glabrous or with sparse to dense J17(15) Body black, broadly oval, strongly convex;
pubescence, rarely forming patterns (with patterns in elytra with tufts of short pubescence in rows; legs
Trogossitidae, some Nitidulidae); lower surface with- retractable into ventral body cavities; scutellum large,
out silky pubescence; elytra with or without striae; triangular ... (Fig. 172) .. [33] NOSODENDRIDAE
scutellary striole absent ... 10 —Body often bicoloured, narrowly oval, moderately
convex to slightly flattened, glabrous; legs not
J10(9) Antennal club 5-segmented; body flat, strongly retractable into ventral body cavities; scutellum small
glossy; elytra with longitudinal striae ... (Fig. 200) .. [52] EROTYLIDAE
... (Fig. 119) .. [9] AGYRTIDAE
—Antennal club 1-3-segmented ... 11 J18(14) Elytral shoulders more or less produced ant-
eriorly; elytral suture with a posterior gap (Fig. 93)
J11(10) Antennal club 1- or 2-segmented; anterior pro- created by widened sutural flanges; pronotum with
notal angles produced (Fig. 185, 189) ... 12 indistinct and incomplete lateral carinae; frons usually
—Antennal club 3-segmented; anterior pronotal angles with a pair of cavities; basal antennal segment enlar-
not produced ... 13 ged, as long as the 2 following segments combined
... (Fig. 197) .. [49] CAVOGNATHIDAE
J12(11) Body elongate-oval; pronotum horseshoe shaped, —Elytral shoulders not produced anteriorly; elytral
with anterolateral angles strongly produced anteriorly; suture complete (Fig. 94) (exception: Cryptophagidae);
pygidium covered by elytra; tarsi pubescent pronotum usually with complete lateral carinae; frons
... (Fig. 185) .. [40] PHYCOSECIDAE without cavities; basal antennal segment slightly en-
—Body narrowly elongate; pronotum oval or rectangu- larged, usually shorter than segments 2+3 ... 19
lar, with anterolateral angles more or less produced;
pygidium exposed; tarsi glabrous; head with promi- J19(18) Body narrowly subparallel, subcylindrical to
nent temples ... (Fig. 188, 189) .. [43] ΜΟΝΟΤΟΜΙDΑΕ slightly flattened, moderately elongate; eyes coarsely
(Lenax, Monotoma) faceted; head, pronotum, and elytra nearly equal in
width; elytra striate or seriate-punctate; tarsal seg-
J13(11) Body ant-like; antennal club loose; dorsal sur- ments 2 and 3 with setose lobes
face usually with long pubescence; head with a distinct ... (Fig. 199) .. [51] LANGURIIDAE
neck, nearly as wide as pronotum; pronotum and elytra — Body broadly to narrowly oval, rarely subparallel; eyes
oval, with humeri rounded finely to moderately coarsely faceted; head distinctly
... (Fig. 122) .. [11] SCYDMAENIDAE narrower than pronotum; elytra as broad as pronotum,
—Body not ant-like; antennal club distinct, usually com- usually without distinct striae; tarsal segments 2 and 3
pact; dorsal surface glabrous or variably pubescent...14 without setose lobes ... 20

-103--
103-
J20(19) Antennae usually closely approximated at base, 3-3-3; antennae 10- or 11-segmented, incrassate or with a
often diverging to form a distinct V; pronotum often l-3-segmented club), Mycetophagidae (tarsi 3-4-4 or 4-4-
depressed at base; body often with silky pubescence 4; antennae 11-segmented with a 3- or 4-segmented club),
... (Fig. 198) .. [50] CRYPTOPHAGIDAE Prostomidae (tarsi 4 4 4; antennae 11-segmented with a 3-
—Antennal bases broadly separated, and antennae not segmented club).
forming a V-shape; pronotum not depressed; body
with short pubescence and sometimes with additional Κ1Tarslfomu2-;bdyinte,oflsha1
longer hairs ... 21 mm long, broadly to narrowly oval; elytra often ex-
posing 1 or 2 abdominal tergites; hind wings feather-
J21(20) Antennal club abruptly compact or ball shaped; like, fringed with long hairs; antennae thin, bearing
tarsal segments 1-3 more or less dilated, and segment long hairs and with a 2- or 3-segmented club
4 sometimes smaller ... (Fig. 118) .. [8] PTINIDAE
... (Fig. 187).. [42] NITIDULIDAE (in part) —Tarsal formula 3-3-3, 3-4-4, or 4-4-4; body usually
—Antennal club loosely formed, not ball shaped; tarsal over 1 mm long, variable in shape; elytra covering
segments simple ... 22 abdomen (exception: Rhizophaginae); hind wings not
featherlike, not fringed with hairs ... 2
J22(21) Body regularly oval and convex or elongate and
Κ2(1) Body broadly oval to hemispherical; head more or
subparallel; pronotal base usually not sinuate; pub-
less concealed by pronotum ... 3
escence short or long, with bristles or scales often
—Body oblong oval to narrowly elongate and sub-
forming patterns
parallel; head not concealed by pronotum ... 5
... (Fig. 179-181) .. [37] TROGOSSITIDAE
—Body irregularly oval and flattened, narrowed post- Κ3(2) Tarsi 4-4-4 segmented, with all segments simple;
eriorly; pronotal base sinuate; pubescence sparse and antennae 8-10-segmented with a 2-segmented club;
short, grouped into clumps on elytra metacoxae expanded into large plates concealing fem-
... (Fig. 191) .. [44] PHLOEOSTICHIDAE (Priasilpha) ora; eyes partly or entirely divided; body small (0.9-
1.5 mm), in life capable of rolling into a ball
... (Fig. 156) .. [18] CLAMBIDAE
Group Κ —Tarsi truly 3-3-3-segmented or appearing 3-3-3-seg-
Includes families with tarsal formula 2-2-2, 3-3-3, 3-4-4, mented, with at least the 2nd bilobed; antennae 7-11-
or 4-4-4; antennae often with a 1-3-segmented distinct segmented with a 1-6- (most often 3)-segmented club;
club; mostly small beetles averaging 2.5-3.0 mm, though metacoxae without large plates; eyes not divided ... 4
some species up to 17 mm long.
Κ4(3) Head partially or entirely concealed by pronotum,
BOSTRICHOIDEA: Jacobsoniidae (see also Group J) (tarsi 2- but anterior part of pronotum usually emarginate and
2-2, or 5-5-5 but appearing 3-3-3; antennae 10-or 11-seg- not extended into a sharp margin over head; body
mented with a 1-3-segmented club). BYRRHOIDEA: Hetero- often larger (1-7 mm), often spotted, usually glabrous
ceridae (tarsi 4 4 4; antennae 11-segmented with a 6- or 7- or subglabrous; antennae shorter than pronotum, with
segmented club). CUCUJOIDEA: Bothrideridae (tarsi 4-4-4; a 1-6-segmented club; maxillary palps securiform or
antennae 11-segmented with a 2- or 3-segmented club), with apex obliquely truncate (Fig. 13-15)
Cerylonidae (tarsi 3-3-3 or 4 4 4; antennae 11-segmented ... (Fig. 206) .. [56] COCCINELLIDAE
with a l- or 2-segmented club), Coccinellidae (tarsi 3-3-3 —Head partially or entirely concealed by pronotum, with
or 4-4-4; antennae 7-11-segmented, incrassate or with a anterior part of pronotum extending into a sharp
1-6-segmented club), Corylophidae (tarsi 3-3-3; antennae margin over head; body small (0.5-2.5 mm), usually
11-segmented with a 3-segmented club), Endomychidae unicoloured, densely pubescent; antennae longer than
(incl. Holoparamecinae) (tarsi 3-3-3; antennae 11-seg- pronotum, with a 3-segmented club; maxillary palps
mented with a 2- or 3-segmented club), Corticariidae not as above ... (Fig. 207) .. [57] CORYLOPHIDAE
(=Lathridiidae) (tarsi 3-3-3; antennae 11-segmented with
a 2- or 3-segmented club), Monotomidae (see also Groups Κ5(2) Pygidium exposed; antennae 10-segmented, with a
F and J) (tarsi 4 4 4; antennae 10-segmented with a 1- or 2- l- or 2-segmented club; tarsal formula 4-4-4
segmented club). EUCINETOIDEA: Clambidae (tarsi 4 4 4; ... (Fig. 188, 189) .. [43] MONOTOMIDAE
antennae 8-10-segmented with a 2-segmented club). (Lenax, Monotoma)
STAPHYLINOIDEA: Ptiliidae (tarsi 2-2-2; antennae 10- or 11- —Pygidium covered by elytra or (Prostomidae) partially
segmented with a2- or 3-segmented club). TENEBRIONOIDEA: exposed; antennae usually 1 1-segmented (10-segmen-
Ciidae (tarsi 4 4 4 or 3-3-3; antennae 8-10-segmented ted in some Colydiidae), with a 1-4-segmented club;
with a 2- or 3-segmented club), Colydiidae (tarsi 4-4-4 or tarsal formula 3-3-3, 3-4-4, or 4-4-4 ... 6

-104-
Κ6(5) Antennae as long as width of clypeus, with a —Body elongate to broadly oval, flattened or moderately
pectinate 7-segmented club; tibiae flattened, spin-ose; convex, rarely cylindrical (exception: some
body robust; elytra subparallel, with paler longi- Colydiidae), variably pubescent, rarely with bristles;
tudinal spots ... (Fig. 162) .. [24] ΗΕΤEROCΕR1DΑΕ head and pronotum without horns; head usually not
—Antennae much longer than width of clypeus, with a 1- deflexed ... 14
4-segmented club; tibiae not spinose; body usually not
robust; elytra rarely with spots ... 7 Κ14(13) Antennal insertions exposed; antennal club f-
or 2-segmented; body usually narrowly elongate and
Κ7(6) Antennae with a loose club; temples strongly pro- slightly flattened, with dorsal surface glabrous and
duced posteriorly; eyes prominent, projecting; body glossy, or with sparse short bristles; elytra striate
strongly depressed, subparallel, 6-8 mm long ... ... (Fig. 203) .. [54] CERYLONIDAE
... (Fig. 222) .. [69] PROSTOMIDAE —Antennal insertions concealed from above; body with
—Antennae with a strongly defined, usually compact dorsal surface usually pubescent ... 15
club; temples not produced posteriorly; eyes moder-
ately to strongly prominent; body convex to slightly Κ15(14) Tarsal formula 3-4-4 (male) or 4-4-4; sides of
depressed, subparallel to broadly oval ... 8 pronotum smoothly continuous with sides of elytra;
pronotum often with 2 depressions near base; body
Κ8(7) Pronotum distinctly narrower than elytral base broadly to narrowly oval, densely pubescent, often
(exception: Lithostygnus, Corticariidae) ... 9 with orange or yellowish spots on elytra or uniformly
—Pronotum as broad as elytral base or slightly narrower brown; antennae usually with a loose 3- or 4-seg-
... 12 mented club; antennal insertions exposed
... (Fig. 209) .. [59] MYCETOPHAGIDAE
Κ9(8) Tarsal formula 4-4-4; antennal insertions exposed; —Tarsal formula 3-3-3 or 4-4-4 in both sexes; sides of
antennae with a compact 2-segmented club; pronotum pronotum not as above; pronotum without 2 basal
usually with a medial impression; elytra with distinct depressions and differently sculptured, often covered
striae; body narrowly oval, depressed, glossy with pubescence or scales forming patterns; antennae
... (Fig. 201, 202) .. [53] BOTHRIDERIDAE with an abrupt, 1-3-segmented club; antennal inser-
—Tarsal formula 3-3-3 ... 10 tions concealed ... (Fig. 215, 216) .. [65] COLYDIIDAE

Κ10(9) Pronotum distinctly waisted, with a basal series

of laterally elongate foveae; antennae with a 2-seg-
mented spherical club and with segments appearing to
be fused together; small beetles, 1.4-1.7 mm long
... (Fig. 205) .. [55] ENDOMYCHIDAE
(Holoparamecinae)
—Pronotum not as above ... 11

Κ11(10) Body less than 0.6 mm long, rust-brown; head

abruptly constricted behind eyes; antennae with club
appearing 1-segmented ... [32] JACOBSONIIDAE
(Derolathus) (see also Group J)
Body 0.8-3.0 mm long, brown to black; head not
abruptly constricted behind eyes; antennae with club
2- or 3-segmented ... (Fig. 208) .. [58] CORTICARIIDAE
(= Lathridiidae)

Κ12(8) Pronotum with a conspicuous raised, sharp carina

on either side of disc (exception: Holoparamecinae)
... (Fig. 204) .. [55] ENDOMYCHIDAE
—Pronotum not as above ... 13

Κ13(12) Body elongate, usually cylindrical, glabrous or

with erect bristles; pronotum and/or head often bear-
ing tubercles or horns; head deflexed; antennal club
large, loose, 2- or 3-segmented; body length 1-3 mm
... (Fig. 211) .. [61] CIDAE

— 105 —
 II: LINEAR KEY 8(5) Tarsi pseudotetramerous (5-5-5 but appearing 4-4-
[provided by J.C. Watt] 4), with segment 3 usually strongly bilobed, rarely
Families with tarsal formula 5-5-5 in females but 5-5-4 in some or pseudotrimerous (4-4-4 but appearing 3-3-3) (Fig.
all males—MONOTOMIDAE, CUCUJIDAE, LAEMOPHLOEIDAE, 67-81) ... CHRYSOMELOIDEA, CURCULIONOIDEA,
CRYPTOPHAGIDAE (all N.Z. genera), PHLOEOSTICHIDAE (all N.Z.
some BYRRHOIDEA.. 9
genera), and PHALACRIDAE (Cy claxyra) —key out in two places. —Tarsi not so ... 19

1 Metacoxae large, fused to metasternum, completely div- 9(8) Antennae lamellate (Fig. 22); pronotum humped
iding 1st ventrite (Fig. 88, 89); sternites 1-3 fused; laterally ... (Fig. 164) .. [26] CHELONARIIDAE
prothorax with distinct notopleural sutures (Fig. 2) —Antennae and pronotum not so ... 10
... ADEPHAGA .. 2
—Metacoxae varying in size, usually movably articu- 10(9) Head without a rostrum, or rarely (Acanthoscelides)
lated to metasternum, never completely dividing 1st slightly rostrate; antennae without a club, not geni-
ventrite (Fig. 90); all sternites usually free; prothorax culate; antennal scrobes absent; gular sutures distinct
without notopleural sutures (Fig. 3) .. POLYPHAGA .. 5 and separate ... CHRYSOMELOIDEA .. 11
—Head usually produced into a rostrum (Fig. 6, 7)
2(1) Hind legs without swimming hairs; prosternum not (rostrum short and not apparent in Scolytinae, Platy-
prolonged behind as a median keel; terrestrial beetles podinae); antennae with a more or less distinct club,
not streamlined for aquatic life ... 3 often geniculate and retractable into scrobes on side of
—Hind legs with swimming hairs; prosternum prolonged rostrum (Fig. 6, d); gular sutures usually confluent
behind as a median keel; body streamlined for aquatic (Fig. 7) or obsolete ... CURCULIONOIDEA.. 13
life ... 4
11(10) Antennae inserted on tubercles, and capable of
3(2) Antennae moniliform (Fig. 19); head and pronotum being flexed backwards against body, usually extend-
with deep, paired, longitudinal grooves ing at least to base of elytra
... (Fig. 101) .. [1] RHYSODIDAE ... (Fig. 234-237) .. [76] CERAMBYCIDAE
—Antennae usually filiform (Fig. 1), never moniliform; —Antennae not inserted on tubercles, not capable of
head and pronotum without deep, paired, longitudinal being flexed backwards against body, usually not
grooves ... (Fig. 102-107) .. [2] CARABIDAE extending to base of elytra ... 12
(incl. Cicindelinae)
12(11) Head somewhat rostrate; antennae and body
4(2) Eyes not divided, or absent (e.g., Kuschelydrus);
bearing scales ... (Fig. 238) .. [77] CHRYSOMELIDAE
hind legs longer than forelegs or middle legs
(Bruchinae)
... (Fig. 108, 109) .. [3] DYTISCIDAE
—Head not at all rostrate; antennae and body without
—Eyes completely divided into upper and lower parts;
scales ... (Fig. 239-243) .. [77] CHRYSOMELIDAE
forelegs distinctly longer than middle or hind legs
(excl. Bruchinae)
... (Fig. 110) .. [4] GYRINIDAE
13(10) Antennae usually geniculate (Fig. 6, 28), with 1st
5(1) Antennae with 10 or fewer segments, the terminal segment retractable into scrobes (Fig. 7, d)
segments lamellate, produced anteriorly into laterally ... (Fig. 251-260) .. [82] CURCULIONIDAE
flattened plates (Fig. 30, 31) ... SCARABAEOIDEA .. 6 —Antennae not geniculate ... 14
—Antennae rarely lamellate or flabellate, if so then 11-
segmented ... 8 14(13) Tarsi appearing 3-3-3; body form characteristic,
as figured .. (Fig. 248) .. [80] BELIDAE (Aglycyderinae)
6(5) Antennae geniculate, with club segments not able to —Tarsi appearing 4 4body
-4; form not so ... 15
be folded closely together; mandibles prominent
... (Fig. 139-143) .. [13] LUCANIDAE 15(14) Labrum distinct, separate (Fig. 6, b) ... 16
—Antennae not geniculate, with club segments able to be —Labrum not distinct, not separate ... 17
folded closely together; mandibles not prominent ... 7
16(15) Prothorax without transverse or lateral carinae;
7(6) Abdomen with 5 ventrites; elytral sculpture very abdominal ventrites free and not strongly sclerotised
rough ... (Fig. 144) .. [14] TROGIDAE ... (Fig. 244) .. [78] NEMONYCHIDAE
—Abdomen with 6 ventrites; elytra smooth, or if —Prothorax with a transverse carina and often additional
sculptured then not rough lateral carinae; ventrites 1-4 fused, strongly sclero-
... (Fig. 145-152) .. [15] SCARABAEIDAE tised ... (Fig. 245, 246) .. [79] ANTHRIBIDAE

—106—


17(15) Body strongly elongate, at least 18 mm long in- abdominal ventrites free; body form not as above; hind
cluding rostrum (Fig. 249) [81] BRENTIDAE
... angles of pronotum at most rectangular, not produced
(Brentinae) backwards 24
...

—Body length considerably less than 18 mm 18...

23(22) Labrum not visible externally; antennal insertions

18(17) Trochanters strongly oblique, not separating fem- distant from eyes (Fig. 165) .. [27] EUCNEMIDAE
...

ora and coxae (Fig. 62); 2 gular sutures, distinct and —Labrum free, visible externally; antennae inserted near
separate (Fig. 247) .. [80] BELIDAE (Belinae)
... eyes (Fig. 166, 167) .. [28] ELATERIDAE
...

—Trochanters slightly oblique, distinctly separating

femora and coxae (Fig. 63); 1 gular suture, positioned 24(22) Minute beetles less than 1 mm long; antennae
medially (Fig. 250) .. [81] BRENTIDAE (Apioninae) slender, filiform, with last 3 segments enlarged; wings
reduced to a central axis with anterior and posterior
fringes of long hairs (Fig. 118) .. [8] PTILIIDAE
...

19(8) Antennae geniculate, with a compact 3-segmented —Larger beetles, length exceeding 1 mm; antennae not
club; elytra truncate, exposing last 2 broad abdominal so; wings (if present) with a broad lamina and venation
tergites, never regularly 9-or 10-striate; body strongly ... 25
sclerotised, usually black, distinctive in form
(Fig. 114, 115) .. [6] HISTERIDAE 25(24) Elytra truncate, leaving usually 6 sternites ex-
—Antennae rarely geniculate, or if so then without a posed (Fig. 124, 127-130) .. [12] STAPHYLINIDAE
...

compact and 3-segmented club; other characters never (in part)

all present in combination ...20 —Elytra rarely truncate, leaving fewer than 6 sternites
exposed (usually 1-3) 26
...

20(19) Metacoxae with posterior face vertical and at least

slightly, usually strongly, excavate to receive retracted 26(25) Loosely articulated, elongate beetles with soft
femur (Fig. 66); antennae filiform, serrate, pectinate, integument (e.g, Fig. 169); elytra not closely em-
or thickened but never with a true club; ocelli absent; bracing abdomen laterally; metacoxae very short,
procoxal cavities open behind (Fig. 40) 21 ... without plate-like extensions of ventral hind margin
—If metacoxae with posterior face vertical and excavate (femoral plates) 27
...

(Nosodendridae, Dermestidae, Anobiidae) then anten- —Compact, usually less elongate beetles, with firmer
nae with a distinct 3-segmented club, or head with a integument; elytra closely embracing abdomen later-
median ocellus, or prothorax produced over head in ally; metacoxae usually longer, and with distinct
the form of a hood; procoxal cavities usually partly or femoral plates (Fig. 52, 53) 28
...

completely closed behind (Fig. 41) 37 ...

27(26) Elytra with longitudinal costae between pairs of

21(20) Abdomen with 2 basal ventrites fused together, striae; trochanters long, transversely joined to femora
the suture partly obliterated; tarsi with ventral adhes- (Fig. 64) (Fig. 168) .. [29] LYCIDAE
...

ive lobes on segments 2-4; body form distinct, cylin- —Elytra without distinct striae or longitudinal costae;
drical anteriorly (Fig. 157) .. [33] BUPRESTIDAE
... trochanters short, obliquely joined to femora (Fig. 65)
—Abdomen with all ventrites usually free, or if fused (Fig. 169) .. [30] CANTHARIDAE
...

(Elateroidea) then suture between 1st and 2nd vent-

rites as distinct as that between 2nd and 3rd; tarsi 28(26) Base of pronotum crenulate; scutellum heart
rarely with adhesive lobes on more than 1 segment; shaped (Fig. 163) .. [25] PTILODACTYLIDAE
...

body form not as above 22

... —Base of pronotum not crenulate; scutellum not so 29 ...

22(21) Anterior median part of mesosternum deeply and 29(28) Metacoxae at least narrowly separated by a tri-
narrowly excavate, with sides of cavity vertical, angular anterior median projection of basal sternite;
receiving narrow, pointed posterior process of body without sessile scales; anterior part of proste-
prosternum, these together usually forming part of rnum somewhat produced forwards to partly cover
`click mechanism' (Fig. 54); abdomen with basal 4 ventral Surface of head BYRRH IDEA (in part) .. 30
... O

ventrites fused; body form characteristic, as in Fig. —Metacoxae contiguous, or if narrowly separated
165-167; hind angles of pronotum almost always (Byrrhidae, Curimus) then body clothed dorsally with
produced backwards, partly around elytral shoulders sessile scales; prosternum emarginate anteriorly 34...

ELATEROIDEA.. 23
...

—Anterior median part of mesosternum shallowly and 30(29) Antennae very short, with anterior processes on
broadly excavate, or not excavate at all; posterior last 7 or 8 segments (Fig. 29); body densely setose or
prosternal process absent, or not shaped as above; with very long pubescence (Fig. 159, 162) 31
...

—107—
 —Antennae not as above; body neither densely setose —Antennae not as above, longer than maxillary palps;
nor with very long pubescence ... 32 head without a Y-shaped impressed line ... 39

31(30) Legs without spines; tarsal formula 5-5-5 38(37) Body usually very convex, elongate-oval to
... (Fig. 159) .. [21] DRYOPIDAE almost hemispherical; abdomen with 5 ventrites
—Legs, especially forelegs, with a row of spines on ... (Fig. 111-113) .. [5] HYDROPHILIDAE
external edge; tarsal formula 4-4-4 —Body less convex, more elongate; abdomen with 6 or 7
... (Fig. 162) .. [24] ΗΕTROCΙDΑ ventrites ... (Fig. 116, 117) .. [7] HYDRAENIDAE
32(30) Metacoxae widely separated, oval; body less than
1.4 mm long; intertidal ... [23] LIMNICHIDAE 39(37) Elytra truncate at apex, leaving at least 3 sclero-
(Hyphalinae) tised abdominal tergites uncovered; antennae filiform
—Metacoxae narrowly separated, transverse; body or thickened towards apex, but without a strong,
larger, length more than 1.4 mm; not intertidal ... 33 compact club (exception: Staphylinidae : Euaesthet-
inae, Microsilphinae) ... 40
33(32) Body elongate, with pubescence not forming a —If 3 or more sclerotised tergites exposed then elytra not
pattern; ventral surface without grooves for legs and truncate (Salpingidae : Inopeplinae), or antennae with
antennae ... (Fig. 160) .. [22] ELMIDAE a strong, compact club (Nitidulidae) ... 41
—Body broadly oval, with pubescence forming a pattern;
legs and antennae retractable into grooves on ventral 40(39) Abdomen with very limited dorsoventral flexibility;
surface... (Fig. 161).. [23] LIMNICHIDAE (Limnichinae) maxillary palps usually long and modified (Fig. 16);
integument with characteristic deep foveae in various
34(29) Metacoxae scarcely contiguous, occasionally not positions, especially on vertex of head and pronotum
contiguous; body broad, convex; mesocoxae widely ... (Fig. 126) .. [12] STAPHYLINIDAE (Pselaphinae)
separated (Fig. 52, 53); legs and antennae retractable —Abdomen flexible dorsoventrally; maxillary palps
into grooves or depressions on ventral surface usually moderately long and not modified; integument
... (FIg. 158) .. [20] BYRRHIDAE rarely with such foveae
—Metacoxae strongly contiguous; body form variable, ... (e.g., Fig. 127-138) .. [12] STAPHYLINIDAE
not as above; mesocoxae narrowly separated; legs and (excl. Pselaphinae, Silphotelus)
antennae usually not retractable into grooves or de-
pressions on ventral surface ... EUCIΝΕTOIDEA .. 35 41(39) Body less than 1.8 mm long, convex; vertex with
a pair of deep foveae between eyes; sides of frons with
35(34) Metacoxae considerably narrowed laterally; fem- a deep semicircular incision in front of each eye
oral plates small; 4th tarsomere lobed below; head ... (Fig. 125) .. [12] STAPHYLINIDAE (Silphotelus)
with characteristic keels behind and below eyes (Fig. —Head without this combination of foveae and lateral
12) ... (Fig. 153, 154) .. [ 16] SCIRTIDAE incisions ... 42
—Metacoxae not narrowed laterally; femoral plates very
large; tarsi without lobed segments; head without such 42(41) Metacoxae with posterior face vertical and ex-
keels ... 36 cavated to receive retracted femur (Fig. 56) ... 43
—Metacoxae with posterior face not vertical ... 47
36(35) Elongate-oval beetles; head narrow, concealed by
pronotum; tarsal formula 5-5-5 43(42) Head with a pair of ocelli between and close to
... (Fig. 155) .. [17] EUCINTDAE eyes; small beetles, length less than 1.8 mm
—Hemispherical beetles; head very broad, reflexible ... (Fig. 170) .. [31] DERODONTIDAE
against a cavity (bounded posteriorly by a semicircular —Paired ocelli absent ... 44
carina) on underside of thorax; tarsal formula 4-4-4
... (Fig. 156) .. [18] CLΑΜΒΙDAE 44(43) Head with a median ocellus (sometimes obscured
by long, coarse setae) ... [34] DERMESTIDAE (in part)
—Head without any ocellus ... 45
37(20) Antennae short, with 6th segment modified as a
cupule (Fig. 26, 27) and terminal 3 or 4 segments 45(44) Prothorax produced over head in a hood-like
forming a strong, pubescent club (exception: Meropa- fashion; antennae serrate, flabellate, or with 3 seg-
thus, with club elongate and not as above — Fig. 117), ments each enlarged but not forming a compact club
or, if club weak and not pubescent, then maxillary ... (Fig. 177) .. [36] ANOBIIDAE (Anobiinae)
palps much longer than antennae; head often with a Y- —Prothorax not hooded; antennae with a compact club
shaped impressed line on vertex ... 38 ... 46

— 108-
46(45) Form very convex, broadly oval; tibiae flattened, -Basal
-Basal 3 ventrites fused, the sutures between them
expanded towards apex; procoxal process received by shallower and narrower than those between 3rd/4th
median cavity in mesosternum and 4th/5th ventrites (which are freely movable,
... (Fig. 172) .. [33] NOSODENDRIDAE usually with exposed intersegmental membranes);
—Form less convex, elongate; tibiae not flattened nor antennae variable in shape, but very rarely with a 3-
expanded apically; procoxal process absent segmented club; body length usually over 3.5 mm
... [34] DERMESTIDAE (Dermestinae) ... (Fig. 219-221) [68] TENEBRIONIDAE

47(42) Tarsal formula 5-5-4 ... 48 56(54) Procoxal cavities closed behind by lateral expan-
—Tarsal formula not 5-5-4 ... 69 sions of apex of prosternal process (Fig. 39); basal 3 or
4 ventrites fused together ... 57
48(47) Head sharply constricted to a narrow neck, best —Procoxal cavities closed behind by inward extensions
seen in oblique views in most specimens ... 49 of hypopleura (Fig. 38), or not completely closed;
—Head not sharply constricted to a narrow neck ... 53 ventrites usually free, rarely with basal 2 fused
together ... 58
49(48) Apical abdominal tergite produced posteriorly
into a stout spine ... (Fig. 213) .. [63] MORDELLIDAE 57(56) Prosternal intercoxal process broad; basal 3 ven-
—Apical abdominal tergite not produced as a spine ... 50 trites fused together, and 4th and 5th movable; scales
usually present ... (Fig. 217) .. [66] ULODIDAE
50(49) Antennae flabellate; tarsal claws toothed (Fig. 87) (formerly as Zopheridae)
... (Fig. 214) .. [64] RHIPIPHORIDAE —Prosternal process very narrow between coxae; basal 4
—Antennae not flabellate; tarsal claws not toothed ... 51 ventrites fused together, only the 5th movable; scales
usually absent ... (Fig. 218) .. [67] CΗΑLCODRΥΙDAE
51(50) Tibial spurs pubescent (Fig. 85)
... (Fig. 233) .. [75] SCRAPTIIDAE
58(56) Tibial spurs serrate (exception: Dοxοzilora) (Fig.
—Tibial spurs absent ... 52 84) ... (Fig. 212) .. [62] MELANDRYIDAE
—Tibial spurs simple or absent ... 59
52(51) First 2 sternites fused, the suture between them
obliterated, at least medially; tarsi with penultimate
59(58) Penultimate tarsal segment bilobed, extending be-
segment small and simple, antepenultimate segment
neath last segment ... 60
lobed below ... (Fig. 232) .. [74] ADERIDAE
—All tarsal segments simple, without lobes ... 62
—First 2 sternites freely movable, the suture between
them distinct; tarsi with penultimate segment lobed
below, antepenultimate segment simple 60(59) Basal 2 ventrites fused together, and ventrites 3-5
... (Fig. 228-231) .. [73] ANTHICIDAE with visible intersegmental membranes
... (Fig. 229) .. [73] ANTHICIDAE
53(48) Elytra abbreviated, leaving at least 4 sclerotised (Lagrioidinae)
tergites exposed ... (Fig. 227) .. [72] SALPINGIDAE —Ventrites free, the lines between equally distinct... 61
(Inopeplinae)
—Elytra not or only slightly abbreviated, covering 61(60) Elytral epipleura indistinct (Fig. 95), restricted to
abdominal tergites, which except for pygidium are proximal (basal) half of elytra; body form as illustrated
membranous ... 54 ... (Fig. 223) .. [70] OEDEMERIDAE
—Elytral epipleura distinct but narrow (Fig. 96), exten-
54(53) With a shelf-like canthus formed by dorsolateral ding to, or almost to apex of elytra; body form as
margin of frons under which antennae are inserted, the illustrated ... (Fig. 224, 225) .. [71] PYROCHROIDAE
canthus extending back and usually encroaching on
front margin of eye (Fig. 11, f); antennal socket and 62(59) Body usually strongly depressed, usually narrow
scape not visible from above ... 55 and parallel sided (Fig. 193-195) ... 63
—Without a true canthus extending back to eye; antennal —Body not strongly depressed, if broad then not par-
socket and base of scape usually visible ... 56 allel sided ... 65

55(54) Only ventrites 1 and 2 fused, remaining ventrites 63(62) Body pubescent; tarsal segment 2 and/or 3 lobed
movable, with visible intersegmental membranes; ant- below or, if not lobed, then antennal scape at least as
ennae with a distinct, loosely articulated, 3-segmented long as the 3 following segments combined; pronotum
club; body length less than 3.5 mm without sublateral carinae and often with anterior
... (Fig. 210) .. [60] ARCHEOCRYPTICIDAE angles produced... (Fig. 192, 193) .. [45] SILVANIDAE

— 109--
109-
 —Body appearing glabrous; tarsal segments 2 and 3 not -Anterior margin of pronotum almost always convex,
lobed below; antennal scape as long as or shorter than covering head, which is not visible from above; if
the 2 following segments combined; pronotum usually pronotum emarginate anteriorly, leaving head partly
with sublateral carinae (Fig. 195) ... 64 visible (Orthoperus), then femoral lines absent
... (Fig. 207) .. [57] CORYLOPHIDAE
64(63) Pronotum without 2 submarginal carinae
... (Fig. 194) .. [46] CUCUJIDAE 73(70) Body characteristically strongly depressed; head
—Pronotum with 2 submarginal carinae sharply constricted to a distinct neck behind prom-
... (Fig. 195) .. [47] LAEMOPHLOEIDAE inent temples; basal 2 ventrites fused
... (Fig. 222) .. [69] PROSTOMIDAE
65(62) Procoxae strongly, obliquely projecting, contiguous —Body convex to moderately depressed; head without a
or almost so, obscuring intercoxal process; body shape distinct neck; all ventrites free, or basal 3 ventrites
as illustrated ... (Fig. 226, 227) .. [72] SALPINGIDAE fused ... 74
—Procoxae not projecting above level of intercoxal
process, which is clearly visible between them; body 74(73) Trochanters of oblique clavicorn type (Fig. 60, d);
shape not as above ... 66 terminal segment of maxillary palp more slender than
penultimate segment, often needle-like; body form as
66(65) Elytra truncate, leaving pygidium exposed; anten- illustrated ... (Fig. 203) .. [54] CERYLONIDAE
nae apparently 10-segmented (segments 10 and 11 —Trochanters of heteromeroid type (Fig. 58, 59), with
fused, forming a 1-segmented club in males) base of femur extending around trochanter on one side
... (Fig. 189) .. [43] MONOTOMIDAE (Monotoma) to coxa; terminal segment of maxillary palp broader
—Elytra not truncate; pygidium not exposed; antennae than penultimate segment ... 75
11-segmented, with a 3-segmented club ... 67
75(74) Basal 3 ventrites fused, the lines between them
67(66) Form broad, convex, almost hemispherical; body fine and shallow in contrast with the broad, deep
glossy, glabrous, 1.6-2 mm long (males of Cyclaxyra) sutures between ventrites 4 and 5 ... 76
... (Fig. 196) .. [48] PHALACRIDAE (in part) —Ventrites free, the divisions between them equal in
—Form elongate, less convex; body matt to slightly depth and width ... 77
glossy, pubescent ... 68
76(75) Antennae 10-segmented; sides of head and ante-
68(67) Male mesocoxal cavities open laterally, reached rior part of prothorax with deep cavities for retracted
by mesepimera; body form as illustrated; length 2.5-5 antennae (Fig. 8, b) ... [68] TENEBRIONIDAE
mm ... (Fig. 190, 191) .. [44] PHLOEOSTICHIDAE (Phrenapatinae)
—Male mesocoxal cavities closed laterally by meeting of —Antennae 11-segmented; sides of head and prothorax
mesosternum and metasternum, not reached by mes- without cavities ... [65] COLYDIIDAE (Pycnomerinae)
epimera; body form as illustrated; length usually less
than 2.5 mm ... (Fig. 198) .. [50] CRYPTOPHAGIDAE 77(75) Antennae 9- or 10-segmented with a 3-segmented
club (Fig. 9); head deflexed (Fig. 9); prothorax some-
what hooded over head (Fig. 9); lateral margins of
69(47) Tarsal formula 4-4-4, rarely 3-4-4 in males frons explanate above antennal base (Fig. 9), but tan-
(Mycetophagidae) ... 70 gential to top of eye, not forming a true canthus
—Tarsal formula not as above ... 80 ... (Fig. 211) .. [61] CIIDAE
—Antennae 11-segmented, usually with a 3-segmented
70(69) Tarsi with antepenultimate segment and often club; head not deflexed; prothorax sometimes extend-
basal segment lobed below; femoral lines often pres- ing forward over head, but not hooded as in Ciidae;
ent on ventrite 1 (Fig. 92, a) ... 71 lateral margin of frons often forming a canthus in line
—Tarsal segments not lobed; femoral lines absent ... 73 with eye and often encroaching on front margin ... 78

71(70) Body form characteristic, with pronotum bearing 78(77) Mesocoxal cavities open laterally (Fig. 42),
a conspicuous raised, curved, sharp carina on each reached by mesepimera; dorsal body surface smooth,
side... (Fig. 204) .. [55] ENDOMYCHIDAE (Mycetaeinae) pubescent, without scales; male tarsi 3-4-4
—Body and pronotum not as above ... 72 ... (Fig. 209) .. [59] MYCETOPHAGID
—Mesocoxal cavities closed laterally (Fig. 43) by meso-
72(71) Anterior margin of pronotum almost always sternum and metasternum, not reached by mesepimera;
emarginate, leaving most of head visible from above dorsal body surface often rough (e.g , Fig. 215) and
.

... (Fig. 206) .. [56] COCCINELLIDAE clothed with scales; tarsi 4-4-4 in both sexes ... 79

-110-
79(78) Body usually subcylindrical, glabrous or micro- smaller than segment 9, 10, or 11, or if antennae
setose, glossy; antennal insertions exposed; all vent- filiform then elytra with transverse striae; protarsi
rites free ... (Fig. 201, 202) .. [53] BOTHRIDERIDAE broader in males than in females; body moderately to
—Body rarely subcylindrical, its surface usually rough strongly convex, oval in outline
and clothed with hairs or scales, usually dull; antennal ... (Fig. 120, 121) .. [ 10] LEIODIDAE
insertions concealed; ventrites 1-3 or 1-4 connate —Antennae not so; elytra without transverse striae;
... (Fig. 215, 216) .. [65] COLYDIIDAE protarsi usually not broader in males than in females
... 88

80(69) Tarsal formula 3-3-3 ... 81 88(87) Antennae weakly but distinctly clubbed, with
—Tarsal formula 5-5-5 ... 84 segments 1-6 glabrous, segment 3 longer than scape,
and segments 7-11 much broader, pubescent, forming
81(80) Body minute, less than 0.7 mm long; antennal a loose club; protarsi and mesotarsi expanded in males
pedicel larger than scape; common in bat guano, and ... (Fig. 119) .. [9] AGYRTIDAE
known from nests of Rattus exulans —Antennae not so; protarsi and mesotarsi rarely
... [32] JACOBSONIIDAE (Derolathrinae) expanded in males ... 89
—Body larger, exceeding 1 mm in length; antennal scape
larger than pedicel ... 82 89(88) Body small, less than 2.5 mm long, glossy, very
convex; elytra truncate apically, exposing sclerotised
82(81) Eyes absent; body parallel-sided pygidium; ventrite 1 at least as long as next 3 ventrites
... (Fig. 201) .. [53] BOTHRIDERIDAE (Anommatinae) together ... (Fig. 131) .. [12] STAPHYLINIDAE
—Eyes present; body not as above ... 83 (Scaphidiinae)
—Body usually larger, if small then not glossy and
strongly convex; elytra not truncate; pygidium not
83(82) Procoxal cavities open behind; body form char- sclerotised; ventrite 1 shorter than next 3 ventrites
acteristic, as illustrated together ... 90
... (Fig. 205) .. [55] ENDOMYCHIDAE (Holoparamecinae)
—Procoxal cavities closed behind; body form not so 90(89) Body shape characteristically ant-like; head
... (Fig. 208) .. [58] CORTICARIIDAE (= Lathridiidae) strongly constricted to a narrow neck; prothorax and
elytra narrowed at base, forming a distinct waist;
84(80) Metasternum longer than combined length of ven- antennae not clubbed; body strongly pubescent;
trites 1-4; legs very short; body depressed, character- terminal segment of maxillary palp short and sharply
istic in shape, as illustrated pointed ... (Fig. 122) .. [11] SCYDMAENIDAE
... (Fig. 171) .. [32] JACOBSONIIDAE (in part) —Body shape not so; other characters not present in
—Metasternum shorter than combined length of ven- combination ... 91
trites 1-4; legs longer; body shape not so ... 85
91(90) Abdomen with pygidium not exposed; antennae
85(84) Prothorax hooded (Fig. 10), produced over head; with last 2 segments fused, forming a conspicuous
head partly or entirely concealed from above ... 86 club; body moderately to strongly elongate
—Prothorax not hooded; head not concealed ... 87 ... (Fig. 188) .. [43] MONOTOMIDAE
(Lenax)
86(85) Antennal insertions separated by more than length —If abdomen with pygidium exposed and sclerotised
of scape; trochanters obliquely joined to femora (Fig. then antennal club 3-segmented and body less
60, a); elytra elongate, parallel-sided elongate ... 92
... (Fig. 174-176) .. [35] BOSTRICHIDAE
(incl. Lyctinae) 92(91) Antennae with a very broad, compact, 3-segmen-
—Antennal insertions closely approximated on frons, ted club; pygidium, and sometimes 1 or 2 tergites in
separated by much less than length of scape; troch- front of it, usually sclerotised and exposed
anters squarely joined to femora (Fig. 61, d); elytra ... (Fig. 187) .. [42] NITIDULIDAE
broad, convex ... (Fig. 178) .. [36] ΑΝΟΒΙΙDΑΕ —Antennae with club (if any) narrower and more loosely
(Ptininae) articulated; abdominal tergites concealed under elytra
... 93
87(85) Antennae with last 5 segments distinctly broader
than basal segments, forming a loosely articulated 93(92) Procoxae strongly projecting and contiguous
club, with segment 8 smaller than segment 7 or 9 (Fig. externally, their cavities oblique; body with stout,
24), or rarely with a 4-segmented club with segment 8 erect pubescence (Fig. 182-184) ... 94
 —Procoxae not or weakly projecting, not contiguous ex- -Elytra without erect setae on dorsal surface, without
ternally, their cavities not oblique; body if pubescent definite rows of punctures; body shape not so; lateral
dorsally then with pubescence shorter, finer, and less margins of pronotum and elytra usually not explanate
projecting, or with short bristles ... 96 ... 103

94(93) Tarsi with at least segments 2 and 3 lobed below 102(101) Protibia with a single spur; antennal club
(exception: Metaxina, all segments simple); antennae usually markedly asymmetrical
usually with a loosely articulated, 3-segmented club ... (Fig. 180) .. [37] TROGOSSITIDAE (Lophocaterinae)
... (Fig. 183, 184) .. [39] CLERIDAE —Protibia with 2 unequal spurs; antennal club sym-
—Tarsi with all segments simple; antennae filiform ... 95 metrical ... [37] TROGOSSITIDAE (Protopeltinae)

95(94) Body bearing very long, erect dorsal setae; mouth- 103(101) Body small, strongly convex, glossy, less than
parts, especially mandibles, prominent; body shape 2.5 mm long; elytral epipleura each with 2 very deep,
characteristic, as illustrated interconnected pits in females
... (Fig. 182) .. [38] CHAETOSOMATIDAE ... (Fig. 196) .. [48] PHALACRIDAE (genus Cyclaxyra)
—Not as above ... (Fig. 186) .. [41] MELYRIDAE —Body not so, more than 2.5 mm long; elytral epipleura
(also as Dasytidae) without deep pits ... 104

96(93) Antennae 10-segmented, with a strong, apparently 104(103) Body broadly oval; sides of pronotum explan-
1-segmented club formed by fusion of segments 10 ate; antennae with last 3 segments much broader than
and 11; body shape characteristic, as illustrated the remainder in females
... (FIg. 185) .. [40] PHYCOSECIDAE ... (Fig. 191) .. [44] PHLOEOSTICHIDAE (Priasilphinae)
—Antennae not as above, usually 11-segmented; body —Body elongate, parallel-sided; sides of pronotum not at
shape not so ... 97 all explanate; antennae with last 3 segments scarcely
differentiated from the remainder ... 105
97(96) Mesocoxal cavities open laterally (Fig. 44), i.e.,
mesepimeron reaching cavity ... 98 105(104) Prothorax with lateral gibbosities, but not car-
—Mesocoxal cavities closed laterally (Fig. 45) by inate laterally in females; body form as illustrated
meeting of mesosternum and metasternum, which ... (Fig. 190) .. [47] PHLOEOSTICHIDAE (Agapythinae)
—Prothorax without lateral gibbosities, carinate later-
exclude mesepimeron from cavity ... 106
ally, at least at apex and base; body form as illustrated
... (Fig. 197) .. [49] CAVOGNATHIDAE
98(97) Body elongate, strongly depressed
... (Fig. 194) .. [46] CUCUJIDAE
—Body usually less elongate, less depressed, and if 106(97) Elytral epipleura indistinct or well marked only
elongate then strongly convex ... 99 in proximal half; epipleural carinae indistinct or
absent posteriorly; body form as illustrated
99(98) Protibia with outer edge characteristic, carinate, ... (Fig. 198) .. [50] CRYPTOPHAGIDAE (females)
with regularly spaced narrow, transverse indentations —Elytral epipleura distinct, extending to apex or almost
(Fig. 83); body usually very small and convex, less so; epipleural carinae distinct throughout ... 107
than 1.7 mm long ... (Fig. 179) .. [37] TROGOSSITIDAE
(Rentoniinae) 107(106) Tarsal claws strongly toothed at base; body
—Protibia with outer edge not so, not carinate; body broad, strongly convex
length exceeding 1.7 mm ... 100 ... (Fig. 196) .. [48] PHALACRIDAE
—Tarsal claws simple; body less convex, more elongate
100(99) Procoxal cavities closed behind; elytra distinctly ... 108
striate ... (Fig. 181) .. [37] TROGOSSITIDAE
(Trogossitinae) 108(107) Pronotum with distinct paired impressions
—Procoxal cavities open behind; elytra without distinct basally; procoxal cavities open behind
striae, though occasionally with numerous rows of ... (Fig. 199) .. [51] LANGURIIDAE
punctures ... 101 —Pronotum without paired impressions basally; pro-
coxal cavities closed behind
101(100) Elytra with numerous long, erect setae on dorsal ... (Fig. 200) .. [52] EROTYLIDAE
surface (Fig. 180), or with numerous rows of punc-
tures; body broad, parallel-sided; lateral margins of
pronotum and elytra explanate ... 102

—112-
 ILLUSTRATIONS

flagellum
pedicel
scape
antenna
mandibles
labrum
clypeus
HEAD
^^ -- tentorial pit
pretarsus
gena
frontoclypeal suture

protibia

lateral
margin PRONOTUM
profemur

humerus

mesofemur

suture
mesotibia

disc — side of
mesotatsus disc ELYTRA

metafemur
metatibia

metatarsus

(1)
/ claws

Fig. 1 Body divisions in Coleoptera: dorsal view, based on Adephaga: Carabidae

[after Lawrence & Britton 1994].

-113-
 mandibles

maxillary palp (palpus)

maxilla labial palp (palpus)
mentum
galea
HEAD submentum ligula
gena pronotal epipleuron

gular suture pleuron

pleurosternal suture

proepisternum notopleural suture

procoxal cavity prosternum

proepimeron procoxa

mesepisternum trochanter
THORAX mesepimeron prosternal process

mesocoxal cavity elytral epipleuron

mesosternum mesocoxa

metepisternum metasternum
metepimeron first visible sternite
(ventńte 1)
metacoxa
metacoxal cavity

ABDOMEN
ventrites

(2)
transverse metasternal suture

Fig. 2 Body divisions in Coleoptera: ventral view, based on Adephaga: Carabidae

[after Lawrence & Britton 1994].

—114—
 notosternal suture
prosternum
prosternal process
hypomeron
procoxal cavity
(open)
mesosternum
mesepisternum
mesepimeron
mesocoxal cavity
(closed)
elytral epipleuron
metepisternum
metasternal — epipleural carina
suture
metasternum
metacoxal cavity
metepimeron

ventrite 1

(3)

Fig. 3 Body divisions in Coleopteta: ventral view, based on Polyphaga: Cryptophagidae

[after Lawrence & Britton 1994].

-115-
 J•ή•
(4)

gaiea maxillary palp

lacinia \ \\
palpifer

subgalea

stipes

cardo

(5)

Fig. 4, 5 Head, ventral, showing mouthparts, and detail of maxilla, ventral (Creophilus oculatus, Staphylinidae).
Flg. 6, 7 Snoutlike head of Curculionidae: (6) dorsal view (a, mandible; b, epistome; c, snout or rostrum; d, scape; e, funicle;
f, club; g, frons; h, vettex; i, occiput;], epicranium); (7) ventral view (a, mandible; b, mouthparts; c, gular suture; d, antennal
scrobe; e, gena; f, occipital foramen; g, occipital area).

-116W--
116-
 (8)

(10)

(12)

49
(13) (14) (15) (16)

Fig. 8-12 Schematic lateral views of head/prothorax: (8) prognathous — Archaeoglenes costipennis (Tenebrionidae) (a,
anterior part of pronotum; b, antennal grooves; c, hypomeron); (9) hypognathous — Cis zeelandicus (Ciidae) (a, pronotum;
b, complete pronotal carina; c, hypomeron; d, procoxa; e, prosternum); (10) hypognathous — Ptinus speciosus (Anobiidae)
(a, prothorax, b, incomplete pronotal carina); (11) head of Artystona erichsoni ( Tenebrionidae), showing canthus (a,
pronotum; b, complete pronotal carina; c, hypomeron; d, procoxa; e, prosternum; f, canthus); (12) head of Veronatus sp.
(Scirtidae), showing characteristic keel on genal area.
Fig. 13-16 Maxillary palps: (13-15) apex obliquely truncate or securiform — Scymnus, Rodalia, Coccinella ( Coccinellidae);
(16) apex elongate, with peglike extension — Physobryaxis inflata (Staphylinidae: Pselaphinae).

—117—
 (19)
(17) (18) (21)

(22)

(28)
(27)
(25) (26)
(23) (24)

(30) (31) (32) (33)

Fig. 17-33 Types of antennae (pubescence not shown): (17, 18) filiform — 17, Cantharidae; 18, Staphylinidae: Habro-
cerinae; (19) moniliform, Rhysodidae; (20, 21) serrate-20, Ptilodactylidae; 21, Lycidae; (22) pectinate, Chelonariidae; (23-
29) clavate-23, abrupt club of Nitidulidae; 24, gradual club with reduced segment 8 of Leiodidae; 25, more evenly developed
club of Lelodidae; 26, 27, segment preceding club a cupule (c), Hydrophilidae; (28) geniculate, Curculionidae; (29) extended,
Dryopidae; (30, 31) lamellate — 30, Lucanidae; 31, Trogidae; (32, 33) flabellate — 32, Bostrichidae; 33, Rhipiphoridae.

-118-
 ANTERlOR pronotal epipreuron pronotal disc

notopleural suture
pieurosternal suture

propleuron
notopleural suture
ANTERIOR

procoxal
procoxal cavIty
cavity
(open posteriorly)

pleurosternal suture

pronotal epipleuron

(34) (35)

ANTERlOR
notosternal suture

.:hypomeron

procoxal notosternal suture

cavity

procoxal cavIty
(closed posteriorly)
(36) (37)

(38)
(39)

Fig. 34 -37 Division of prothorax in the suborders of Coleoptera, ventral and lateral: (34, 35) Adephaga (Maoripamborus
fairburni, Carabidae); (36, 37) Polyphaga (Mimopeus subcostatus, Tenebrionidae).
Fig. 38, 39 Types of procoxal closure: (38) tenebrionid; (39) zopherid / ulodid (a, prosternum; b, prosternal process; c,
hypomeron; d, coxa; e, trochanter; f, femur; g, concealed membranous extension; h, aperture; i, membrane).

-119-
 prosternum ■
Ι procoxal cavity

procoxal cavity

CLOSED PROCOXAL CAVITY

(40)

OPEN PROCOXAL CAVITIES (41)

mesepimeron

mesosternum
mesocoxal
mesepisternum cavity
mesepimeron
elytral
epipleuron
mesosternum elytral
epipleuron
mesoc oxal
c avity metasternum
metepisternum

metasternum

L metepisternum

OPEN MESOCOXAL CAViTY (43) CLOSED MESOCOXAL CAVITY

(42)

mesocoxal
cavity
mesepisternum

mesepisternum
mesepimeron
mesosternum
mesocoxal mesosternum mesepimeron
cavity

metepisternum

metepisternum

metasternum metasternum

L
1

OPEN MESOCOXAL CAVITY
(44) (45) CLOSED MESOCOXAL CAVITY

Fig. 40-45 Types of coxal closure: (40) procoxae completely open behind, Nascioides enysi (Buprestidae); (41) procoxal
cavity closed, Rygmodus incertus (Hydrophilidae); (42) mesocoxal cavity open laterally, Typhaea stercorea
(Mycetophagidae); (43) mesocoxal cavity closed laterally, Enarsus bakewelli (Colydiidae); (44) mesocoxal cavity open
laterally, Platisus sp. (Cucujidae); (45) mesocoxal cavity closed laterally, Cryptodacne brouni (Erotylidae).

—120-
 end of radial bar
RA 3+4 radIal celI
mp-cua crossvein

oblongum cell
κ
ΑΡ
3+4
a
ω
A medial fleck

(46)
radiat hinge

radial cell

oblongum cell

(48)
medial hinge
medial bar

RA 1+2
SC P
Sc A
ScP

radiaI cell

Mp 1+2
ΜΡ 3a
AP 3+4
ΜΡ 3b
AA 3+4
(47) CuA+AA ΜΡ4

wedge cell

Fig. 46-49 Hind wing venation in Coleoptera: (46) example from Adephaga; (47) example from Polyphaga; (48, 49) position
of radial Ioop (radial cell) and medial loop (oblongum cell) in the unfolded and folded wing in Adephaga, after Kukalová-Peck
& Lawrence (1993). Veins: A, anal; AA, anal anterior; AP, anal posterior; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus
posterior; J, jugal; M, media; MA, media anterior; MP, media posterior; R, radius; RA, radius anterior; RP, radius posterior;
r, radial cross-vein; Sc, subcosta; ScA, subcosta anterior; ScP, subcosta posterior.

-121-
 radial spring

Archostemata apical hinge

RA1. 2

r, ScP\ScP
RA 3
r4
RΑ 3

radial spring

flattened ScP apical hinge

RA

ι4
Adephaga
apical hinge
radial spring

(50 )

Hypothetical Ancestor

r3 r4+RP,+RP 2
Ρ? 1 RP 3 ,,
RP,
Archostemata eιiιαt sP
υ' \ RP 2
Μ,ο

Polyphaga Ά.

medial hinge
RP, RP2 ά,
r4 /l //
ΠΡ3 medial spring
Adephaga Q -

U ,επρ2

RP

RP,
RP2

RP
medial hinge

(51) Hypothetical Ancestor

medial spring

Fig. 50, 51 Evolution of end of radial loop (50) and medial loop (51) in Coleoptera. Veins: Ccros-vein;MP,mdaptro
r, radial cross-vein; RA, radius anterior; RP, radiuspoter;—m1and2,io-elcrsvn;SP,ubota
posterior [Fig. 48-51 after Kukalová-Peck & Lawrence 1993].

-122-
 (54)

(55)

(56)

Fig. 52-56 Structural features of ventral thorax and abdomen: (52) Synorthus insularis (Byrrhidae), after Watt (1971) —
a, antennal groove; b, epipleuron; c, crural or femoral depression; d, prosternal process; e, mesosternal depression for
receiving prosternal process; f, mesosternum; g, metasternum; h, metacoxa; i, ventrite 1; (53) Epichorius longulus (Byr-
rhidae), after Watt (1971)—a, mesepisternum; b, mesepimeron; c, epipleuron; d, metepisternum; e, metepimeron; (54) click
mechanism' in Elateridae — a, prosternal process; b, mesosternal cavity receiving prosternal process; (55) Priaslipha
obscura (Phloeostichidae) — a, antennal groove; b, prosternal process; c, mesepisternum; d, mesepimeron; (56) Dermestes
maculatus, Dermestidae — a, coxa with extended femoral plate; b, trochanter; c, femur; d, femoral excavation; e, meta-
sternum; f, metepisternum; g, metepimeron; h, ventrite 1. Scale lines = 1 mm.

-123—
 trochantin

trochanter ‚- femur

trochanter
(62)

(63)
trochanter

trochanter
y
coxal posterior face

coxal excavation

(64) (66)

(59)

(60) (61)

Fig. 57 Divisions of foreleg (Creophilus oculatus, Staphylinidae). Fig. 58-66 Coxa/femur articulation through ttochanter:
(58) heteromeroid type, oblique ventral, Phymatophaea lugubris (Cleridae) - a, prosternal intercoxal process; b, coxa; c,
trochanter; d, femur; (59) heteromeroid type, Pycnomerus sophorae (Colydiidae) - a, basal part of femur; c, coxa; d,
trochantet; (60) oblique mesotrochanterofemoral junction, Euderia squamosa (Bostrichidae) - a, basal part of femur, d,
trochanter; (61) transverse mesotrochanterofemoral junction, Ptinus speciosus (Anobiidae: Ptininae) - a, basal part of
femur; c, coxa; d, trochanter; (62) femur contiguous with coxa, not separated by trochanter, Agathinus tridens ( Belidae:
Belinae); (63) femur separated from coxa by trochanter, Apion sp. (Brentidae); (64) metatrochanter Iong, transversely joined
to femur, Porrostoma rufipenne (Lycidae); (65) metatrochanter short, obliquely joined to femur, Asilis fumida(Canthride);
(66) metacoxa excavated, with posterior face vertical, Nascioides enysi ( Buprestidae).

—124—
 (68) (69)
(67)

(76)
(73) (75)
(72)

(77)

(78) (79)
Fig. 67-81 Protarsal configuration in Chrysomeloidea and Curculionoidea, 5-5-5 with reduced 4th segment (67-79) and
4-4-4 with reduced 3rd segment (80, 81): (67) Arnomus sp., Chrysomelidae; (68) Acanthoscelides obtectus, Chrysomelidae;
(69) Adoxia sp., Chrysomelidae — 4th and 5th segments often imperceptibly fused; (70) Caccomolpus sp., Chrysomelidae;
(71) Alema spatiosa, Chrysomelidae; (72) Eualema speculifera, Chrysomelidae; (73) Allocharis sp., Chrysomelidae; (74)
Prionoplus reticularis, Cerambycidae; (75) Anagotus fairburni, Curculionidae; (76) Cacephatus incertus, Anthribidae; (77)
Rhinorhynchus rufulus, Nemonychidae; (78) Agathinus tridens, Belidae; (79) Lasiorhynchus barbicornis, Brentidae; (80)
Aralius wollastoni, Belidae; (81) Neocyba metrosideros, Btentidae.
Fig. 82 Right metatarsus, Coccinella leonina ( Cucujoidea:Coccinellidae), showing reduced 3rd segment (shaded).

-125-
 (85)

(83)

(91)

(88) (89) (90)

Fig. 83-87 Diagnostic features of coleopteran legs: (83) protibia with transversely indented outer carina, Rentonidium
costiventris (Trogossitidae: Rentoniinae); (84) right metatarsus and tibial spurs, Hylobia pulla (Melandryidae); (85) basal
metatarsal segment and tibial spur, Nothotelus usitatus (Scraptiidae); (86) serrate mesotarsal claws, Tanychilus sophorae
(Tenebrionidae: Alleculinae) -a, empodium; (87) toothed tarsal claw, Allocinops brookesi ( Rhipiphoridae).
Fig. 88-92 Diagnostic features of coleopteran abdomen: (88, 89)1st ventrite bisected by metacoxae (Adephaga — Lancetes
lancelatus, Dytiscidae and Loxomerus nebrioides, Carabidae); (90) 1st ventrite undivided by metacoxae (Polyphaga —
Platisus sp., Cucujidae); (91) intersegmental membranes (free ventrites), Baeocera scutellaris (Staphylinidae: Scaphidi-
inae); (92) patt metathorax and abdomen, Cocinellidae (a, femoral line; b, ventrite 1; c, epipleuron; d, epipleural carina).

-126-
 (93) (94)

(95) (96)

(97)

(98 ) (100)

Fig. 93-96 Diagnostic features of coleopteran elytra: (93) elytral suture with a posterior gap, Neocercus electus (Cavo-
gnathidae); (94) elytral suture complete, Loberus depressus (Languriidae); (95) elytral epipleura indistinct, Parisopalpus
maclearyi (Oedemeridae); (96) elytral epipleura distinct, Techmessa telephoroides (Pyrochroidae).
Fig. 97-100 Morphological types of beetle larvae: (97) eruciform, Eucolaspis brunnea (Chrysomelidae); (98) scarab-
aeiform, Costelytra zealandica (Scarabaeidae); (99) apodous, Sitona discoidea (Curculionidae); (100) campodeiform,
Arpediomimus kronei (Staphylinidae).

Fig. 101-260 Habitus studies of representative members of the families of New Zealand Coleoptera (individually captioned).
Scale lines 1.0 mm unless otherwise indicated. [Expanded captions commence on p. 168.]

Illustrators: Fig. 1-3, 101-107, 110-118, 120-145,148, 149, 151, 153, 155, 156, 158, 159, 161-165, 168, 169, 171-186,
188-205, 207-214, 216-228, 230-260 D. W. Helmore; Fig. 4-47, 52-100 J. Klimaszewski; Fig. 48-51 J. Kukalová-Peck;
Flg. 108, 114, 119, 138, 146, 152, 154, 157, 160, 166, 170, 187, 206, 215, 219 A. C. Harrls; Fig. 147 S. Forgie; Flg. 150 L.
Alexander; Flg. 167 J. Liddiard. Fig. 109 after Ordish (1976a); Fig. 124, 125, 132 after Steel (1964, 1966, 1950α).
-127-
 E
E

(101) Rhyzodiastes proprius

Rhysodidae (103)Maoripamborus fairburni
Carabidae: Carabinae

(104)Loxomerus nebrioides
Carabidae: Migadopinae

(102) Neocicindela tuberculata

dh ie Carabidae: Cicindelinae

—128—
(105) Clivina basalis
Carabidae: Scaritinae

(107) Ctenognathus novaezelandiae

Carabidae: Harpalinae

(106) Zecillenus alacris (108) Liodessus plicatus

Carabidae: Trechinae Dytiscidae: Hydroporinae
-129 -
 (111) Horelophus walkeri
Hydrophilidae: Horelophinae
(109) Kuschelydrus phreaticus
Dytiscidae: Hydroporinae

(110) Gyrinus convexiusculus (112) Enochrus tritus

Gyrinidae: Gyrininae Hydrophilidae: Hydrophilinae
-130-
 —

(115)
(115) Tomogenius latipes
Histeridae: Saprininae

(113) Rygmodus tibialis

Hydrophilidae: Sphaeridiinae

(114) Reichardtia pedatrix (116) Podaena latipalpis
Histeridae: Saprininae Hydraenidae: Hydraeninae

—131 —
(117) Meropathus zelandicus
(119) `Necrophilus' prolongatus
Hydraenidae: Ochthebiinae Agyrtidae

(118) Notoptenidium lawsoni (120) Inocatops elongellus

Ptiliidae: Ptiliinae Leiodidae: Camiarinae

-132-
 (121) Colon hirtale
Leiodidae: Coloninae (123) Microsilpha litorea
Staphylinidae: Microsilphinae

(122) Adrastia clarkei (124) Omaliomimus albipennis

Scydmaenidae: Scydmaeninae Staphylinidae: Omaliinae
-133 -

(125) Silphotelus nitidus (127) Pseudophloeocharis australis

Staphylinidae: Proteininae Staphylinidae: Phloeocharinae

(126) Sagola laminata (128) Sepedophilus sp.

Staphylinidae: Pselaphinae Staphylinidae: Tachyporinae

-134-

(129) Habrocerus capillaricornis (131) Baeocera scutellaris

Staphylinidae: Habrocerinae Staphylinidae: Scaphidiinae

(130) Aleochara hammondi (132) Parasiagonum hudsoni

Staphylinidae: Aleocharinae Staphylinidae: Plestinae

-135 -

(133) Nototorchus ferrugineus (135) Agnosthaetus vicinus

Staphylinidae: Osoriinae Staphylinidae: Euaesthetinae

(134) Carpelimus sp. (136) Pseudopsis arrow!

Staphylinidae: Oxytelinae Staphylinidae: Pseudopslnae

-136-

(137) Medon zeelandicus (139) Ceratognathus parrianus

Staphylinidae: Paederinae Lucanidae: Aesalinae

(138) Ca fius litoreus (140) Paralissotes reticulatus,

Staphylinidae: Staphylininae male, Lucanidae: Lucaninae

—137 —
s
s

(141) Geodorcus auriculatus, (143) Dendroblax earlii

male, Lucanidae: Lucaninae Lucanidae: Lampriminae

(142) Geodorcus auriculatus, (144) Trox scaber

female, Lucanidae: Lucaninae Trogidae

-138-
(145) Acrossidius tasmaniae
Scarabaeidae: Aphodiinae (147) Onthophagus granulatus
Scarabaeidae: Scarabaeinae

(146) Phycochus graniceps (148) Odontria giveni Watt

Scarabaeidae: Aphodiinae Scarabaeidae: Melolonthinae

-139 -
 (149) Costelytra zealandica
Scarabaeidae: Melolonthinae

(151) Heteronychus arator

Scarabaeidae: Dynastinae

(150) Prodontria lewisi (152) Pericoptus truncatus

Scarabaeidae: Melolonthinae Scarabaeidae: Dynastinae
—140—
(153) Amplectopus pallicornis
(155) Eucinetus stewarti
Scirtidae
Eucinetidae

(154) Veronatus tricostellus (156) Clambus domesticus

Scirtidae Clambidae
—141 —

(157) Nascioides enysi (159) Parnida agrestis

Buprestidae Dryopidae

(158) Liochoria huttoni (160) Hydora picea

Byrrhidae: Byrrhinae Elmidae: Larainae

-142-
 (161) Limnichus nigripes
(163) Byrrocryptus urquharti
Limnichidae: Limnichinae
Ptilodactylidae: Anchytarsinae

(162) Heterocerus novaeselandiae

(164) Brounia thoracica
Heteroceridae: Heterocerinae Chelonariidae

-143 -
 E
E

(165) Neocharis simplex (167) Amychus candezei

Eucnemidae: Melasinae Elateridae: Denticollinae

(166) Thoramus wakefieldi (168) Porrostoma rufipenne

Elateridae: Ag ry pninae Lycidae

—144—
 (169) Asilis fulvithorax (171) Saphophagus minutus

Cantharidae: Dysmorphocerinae Jacobsoniidae

(170) Nothoderodontus gourlayi (172) Nosodendron zealandicum

Derodontidae: Laricobiinae Nosodendridae

—145 —
(173) Trogoderma maestum

(175) Dinoderus minutus

Dermestidae: Megatominae Bostrichidae: Dinoderinae

^φ § ^ί^πλ19^a
.ι ^ ., s >9 ι,

(176) Lyctus brunneus

(174) Euderia squamosa
Bostrichidae: Euderinae Bostrichidae: Lyctinae

— 146 —
 (179) Rentonidium costiventris
Trogossitidae: Rentoniinae
(177) Hadrobregmus magnus
Anobiidae: Anobiinae

(178) Ptinus speciosus (180) Grynoma regularis

Anobiidae: Ptininae Trogossitidae: Lophocaterinae

—147 —

(181) Lepidopteryx nigrosparsa (183) Metaxina ornata

Trogossitidae: Trogossitinae Cleridae: Thaneroclerinae

(182) Chaetosoma scaritides (184) Phymatophaea violacea

Chaetosomatidae Cleridae: Enopliinae

-148-
 (187) Platipidia asperella
Nitidulidae: Nitidulinae
(185) Phycosecis limbata
Phycosecidae

(186) `Dasytes' subcyaneus (188) Lenax mirandus

Melyridae: Dasytinae Monotomidae: Monotominae

-149 -

(189) Monotoma spinicollis (191) Priasilpha obscura
Monotomidae: Monotominae Phloeostichidae: Priasilphinae

(190) Agapytho foveicollis (192) Brontopriscus pleuralis

Phloeostichidae: Agapythinae Silvanidae: Brontinae

-150-

(193) Oryzaephilus surinamensis (195) Microbrontes lineatus

Silvanidae: Silvaninae Laemophloeidae

(196) Cyclaxyra impressa

(194) Platisus sp.
Cucujidae Phalacridae: Cyclaxyrinae

—151 —

(197) Zeonidicola chathamensis (199) Hapalips prolixus

Cavognathidae Languriidae: Xenoscelinae

(198) Thortus ovalis (200) Thallis polita

C ryptophagidae: Cryptophaginae Erotylidae: Dacninae

—152—

(201) Anommatus duodecimstriatus (203) Hypodacnella rubriceps

Bothrideridae: Anommatinae Cerylonidae: Euxestinae

(202) Ascetoderes obsoletus (204) Mycetaea subterranea

Bothrideridae: Bothriderinae Endomychidae: Mycetaeinae

-153 -
 (207) Anisomeristes sharpi
Corylophidae: Sericoderinae
(205) Holoparamecus tenuis
Endomychidae: Holoparamecinae

(206) Coccinella leonina (208) Enicmus caviceps

Coccinellidae: Coccinellinae Corticariidae: Lathrldiinae

-154-
 (209) Triphyllus hispidellus (211) Cis zeelandicus
Mycetophagidae: Mycetophaginae Ciidae: Ciinae

(210) Archeoc rypticus topali (212) Hylobia nubeculosa

Archeocrypticidae Melandryidae: Melandryinae

-155 -
 (215) Pristoderus antarcticus
(213) Mordella antarctica Colydiidae: Colydiinae
Mordellidae: Mordellinae

(214) Rhipistena lugubris (216) Rhizonium antiquum

Rhipiphoridae: Pelecotominae Colydiidae: Colydiinae

-156-

(217) Syrphetodes marginatus (219) Chaerodes trachyscelides

Ulodidae Tenebrionidae: Lagriinae

(218) Chalcodrya variegata (220) Actizeta albata

Chalcodryidae Tenebrionidae: Pimeliinae

—157 —
 (221) Mimopeus elongatus
(223) Thelyphassa lineata

Tenebrionidae: Tenebrioninae
Oedemeridae: Oedemerinae

(222) Dryocora howitti (224) Techmessa concolor

Prostomidae Pyrochroidae: Pilipalpinae

-158-
 (227) Diagrypnodes wakefieldi

(225) Exocalopus pectinatus Salpingidae: Inopeplinae
Pyrochroidae: Pilipalpinae

(226) Salpingus bilunatus (228) Anthicus hesperi

Salpingidae: Salpinginae Anthicidae: Anthicinae

—159 —
 (229) Lagrioida brouni
Anthicidae: Lagrioidinae
(231) Macratria exilis
Anthicidae: Macratriinae

(230) Cotes crispi (232) `Xylophilus' nitidus

Anthicidae: Lemodinae Aderidae

—160—
 (235) Zorion sp.
Cerambycidae: Cerambycinae

(233) Nothotelus usitatus

Scraptiidae

(234) Prionoplus reticularis (236) Oemona hirta

Cerambycidae: Prioninae Cerambycidae: Cerambycinae
—161 —
 (239) Ch rysolina hyperici
Chrysomelidae: Chrysomelinae

(237) Xylotoles costipennis

Cerambycidae: Lamiinae

(238) Bruchidius villosus (240) Agasicles hygrophila

Chrysomelidae: Bruchinae Chrysomelidae: Galerucinae (Alticini)

-162-
 (241) Adoxia vulgaris
Chrysomelidae: Galerucinae
(Galerucini)
(243) Eucolaspis brunnea
Chrysomelidae: Eumolpinae

(242) Arnomus broun (244) Rhinorhynchus rufulus

Chrysomelidae: Ctyptocephalinae Nemonychidae: Rhinorhynchinae

-163 -

(245) Sharpius venustus (247) Cyrotyphus tridens
Anthribidae: Anthribinae Belidae: Belinae

(246) Dysnocryptus pallidus (248) Aralius wollastoni

Anthribidae: Choraginae Belidae: Aglycyderinae

—164—
 (250) Neocyba metrosideros
Brentidae: Apioninae

(249) Lasiorhynchus barbicornis

Brentidae: Brentinae

(252) Mandalotus miricollis (251) Anagotus turbotti

Curcullonidae: Brachycerinae Curculionidae: Brachycerinae
—165 —
 (253) Andracalles spurcus
Curculionidae: Curculioninae
(255) Stephanorhynchus lawsoni
Curculionidae: Curculioninae

E
E

(254) Myrtonymus zelandicus (256) Macroscytalus remotus
Curculionidae: Curculioninae

Curculionidae: Cossoninae

-166-
 (259) Phloeosinus cupressi
(257) Xenocnema spinipes Curculionidae: Scolytinae
Curculionidae: Cossoninae

(258) Hylastes ater (260) Platypus apicalis

Curculionidae: Scolytinae Curculionidae: Platypodinae

-167 -
 EXAMPLES OF SPECIES
[1] Rhysodidae labrum; hind wings absent. Found under fallen branches
Fig. 101 Rhyzodiastes proprius (Broun) and at the base of plants, feeding on other ground-dwelling
A black, glossy, subparallel-sided beetle 6-7 mm long, arthropods. In coastal areas from Port Waikato north, and
living under bark or in wood of partly decayed logs. Known especially common on offshore islands.
from Northland, Auckland, Waikato, and Great Barrier l.
[3] Dytiscidae: Hydroporinae
[2] Carabidae: Cicindelinae Fig. 108 Liodessus plicatus Sharp
Fig. 102 Neocicindela tuberculata (Fabricius) An elongate-oval, slightly depressed, black beetle 2.5-2.7
A conspicuously two-toned tiger beetle 10-12 mm long, mm long. Elytra strongly punctate, Iacking striae; protarsi
mostly dark with metallic green reflections and irregular and mesotarsi expanded in male, and metatarsi with a
lateral cream-coloured areas on elytra. Antennae and legs fringe of swimming hairs. Primarily a pond dweller,
long and slender; eyes Iarge and prominent; elytra with inhabiting alpine tarns at 1700 m as well as lowland and
striae, closely punctate, and with a row of about 4 foveae temporary ponds, and even warm pools in the thermal
near suture; mandibles Iarge and sharp-pointed. Abundant region of Rotorua, and found in quiet reaches of streams.
in dry, open habitats such as clay banks, where it preys on Adults have been obsetved in flight. Recorded from North
otherinsects.Movesveryswiftly,andtakestoflightrapidly. Cape to Stewart I.
Larvae live in burrows in exposed warm areas. Occurs on
both main islands. [3] Dytiscidae: Hydroporinae
Flg. 109 Kuschelydrus phreaticus Ordish
[2] Carabidae: Carabinae A small, subterranean diving beetle 1.5-1.6 mm long,
Fig. 103 Maoripamborus fairburni Brookes narrowly elongate, subparallel-sided, brown. Hind legs and
A Iarge, black ground beetle 20-24 mm long, with slightly body, particularly laterally, with long, erect, fine setae;
iridescent upper body sides. Head distinctly elongate, pronotum with sides arcuate. Collected from wells and
projecting in front of protruding hemispherical eyes; man- bores in the northern part of the South I. [Redrawn after
dibles strong, broad. A predator of snails. Recorded mainly Ordish 1976a]
from Waipoua Forest, Northland.
[4] Gyrinidae
[2] Carabidae: Migadopinae Fig. 110 Gyrinus convexiusculus Macleay
Fig. 104 Loxomerus nebrioides Guérin A black, glossy, boat-shaped whirligig beetle 4.3-4.5 mm
A medium-sized ground beetle 15-17 mm long, reddish long. Antennae short, with segments co-adapted; hind legs
dark brown or black with reddish-brown appendages. paddle-shaped, with a fringe of long swimming hairs.
Pronotum narrowed posteriorly; elytra oval. Recorded from Occurs on the surface of ponds, rowing rapidly over the
the Auckland Is. water and preying on other insects. The divided eye
(arrowed) enables it to see simultaneously above and
[2] Carabidae: Scaritinae below the surface. Most recently collected on the Ahipara
Fig. 105 Clivina basalis Chaudoir Plateau, Northland.
A moderately small, pedunculate ground beetle 6-8 mm
Iong, dark brown to almost black with reddish appendages [5] Hydrophilidae: Horelophinae
and basal half of elytra. A chatacteristic narrow waist Fig. 111 Horelophus walkeri Orchymont
between thorax and elytra. Ranging from central North I. to A weakly convex, elongate beetle 2.0-3.0 mm long. Body
Spirits Bay in the north. outline interrupted between pronotum and elytra; pronotal
sides sinuate; antennae very long, with a loosely formed 3-
[2] Carabidae: Trechinae segmented club. Described from 2 specimens collected at
Flg. 106 Zecillenus alacris (Broun) Reefton, and with subsequent records from Cawthron
A small ground beetle 4.0-5.5 mm long, brown with paler, Park, Nelson, this unique species represents an endemic
yellowish-brown appendages and lateral elytra. Eyes large; New Zealand subfamily. Specimens collected on rocks in
posterior pronotum strongly narrowed; elytra with 2 the spray zone of waterfalls suggest that it may be
posterolateral projections. Collected from under logs on the semiaquatic.
beach and from sand near a stream at Karekare and
Whatipu, near Auckland. [5] Hydrophilidae: Hydrophilinae
Fig. 112 Enochrus tritus (Broun)
[2] Carabidae: Harpalinae An elongate-oval, yellowish-brown beetle 4.4-4.5 mm long.
Flg. 107 Ctenognathus novaezelandiae Fairmaire Maxillary palps longer than antennae; elytra punctate, not
A very agile, elongate, glossy black ground beetle 11-12 striate. An adventive Australian species occurring in New
mm long. Antennae and legs thin; elytra regularly striate, Zealand in small ponds; common in cattle troughs in the
fused; 2 prominent tactile setae above eye and a row on Kermadec Is. Adults and larvae feed on algae.

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[5] Hydrophilidae: Sphaeridiinae [9] Agyrtidae
Fig. 113 Rygmodus tibialis Broun Fig. 119 `Necrophilus' prolongatus
A strongly convex, oval beetle 6.5-8.0 mm long, metallic A moderately Iarge, broadly oval, depressed beetle 9.0-10
black with legs, maxillary palps, and basal antennal mm long, dark brown with sides of pronotum and elytra
segments paler and often with a reddish tinge; moderately yellowish brown, explanate. Protarsi and mesotarsi ex-
to densely punctate. Antennae long, with a loose 3- panded in male; elytra striate; antennae with an indistinct 5-
segmented club. Originally described from Mt Arthur, north- segmented club. Recorded from the North I.
west Nelson; we have seen specimens of presumably this
species from L. Rotorua, Salisbury, and Mt Robert. [10] Leiodidae: Camiarinae
Fig. 120 Inocatops elongellus Broun
[6]
Histeridae: Saprininae A small carrion beetle 2.5-2.9 mm long, reddish with
Fig. 114 Reichardtia pedatrix (Sharp) whitish pubescence. Pronotum narrowed posteriorly, with a
A robust, oblong, strongly convex, glossy black pill beetle trans-verse basal impression; elytra oval in outline;
4.8-5.0 mm long. Elytra abbreviated, exposing last 3 seg- antennae with a 4-segmented club, its 2nd segment
ments of abdomen; legs stout, with numerous spines; front narrower than the others. Found in leaf litter. Recorded
legs well adapted to burrowing. Occurs on sandy beaches, from the Nelson area.
often under dead fish or birds, occasionally under sea-
weed. Recorded from Northland to Stewart I. [10] Leiodidae: Coloninae
Fig. 121 Colon hirtale (Broun)
[6] Histeridae: Saprininae An oblong, moderately convex small carrion beetle 2.3-2.5
Fig. 115 Tomogenius latipes (Broun) mm long, pubescent, dull dark brown, with legs and ant-
An oval, convex, glossy black pill beetle 4.5-4.7 mm long. ennae red; antennal club darker, 4-segmented. Terminal
Antennae stout, bearing a compact club; elytra abbreviated segment of maxillary palps small and sharply pointed; head
apically, leaving pygidium exposed; forelegs flattened for with a distinct neck. Widely distributed, and frequently
digging,with approximately 6 teeth along outer edge, and caught in flight interception traps.
other legs bearing stout bristles. At Omahuta found in a
short-tailed bat roost in a hollow fallen kauri tree, on Great [11] Scydmaenidae: Scydmaeninae
Bar ier Island found in kingfisher nests. Adults and larvae Flg. 122 Adrastia clarkei (Franz)
probably feed on fly larvae in guano. Recorded from An elongate-oval beetle 3.3-3.5 mm long, reddish brown,
Omahuta State Forest to Codfish I. near Stewart I. glossy, sparsely pubescent. Antennae 10-segmented;
apex of elytra not fully covering abdomen; last segment of
[7] Hydraenidae: Hydraeninae maxillary palp small; head with a neck, and a marked con-
Fig. 116 Podaena latipalpis Ordish striction between prothorax and abdomen forming a distinct
A small cascade beetle 1.9 mm long, reddish brown with waist'. Occurs in leaf litter. Recorded mostly from the
paler clypeal border and humeral area of elytra. Maxillary South I. (originally found in the Routeburn Valley) and from
palps long, reaching beneath eyes; elytra with 7th interstria the central North Island (Desert Road).
bearing a low carina. Adults have been collected from
October to June from streams in the North I. [12] Staphylinidae: Microsilphinae
Flg. 123 Microsilpha litorea Broun
[7] Hydraenidae: Ochthebinae A small beetle 2.2-2.9 mm long, dark btown with reddish
Flg. 117 Meropathus zelandicus Ordish tarsi and basosutural part of elytra. Sparsely pubescent;
A uniformly dark brown cascade beetle 2.5 mm long, with elytra covering most of abdomen; protibae strong, flat-
paler, dense, strongly recumbent setae on pronotum and tened, bearing teeth and spines; antennae with club 4-
elytra. Head and thorax with granulose microsculpture; segmented. Known only from the beach at Port Chalmers,
elytra glabrous between microsculpture and setae. Occurs Dunedin, although most other species inhabit forests and
in litter, and is typically covered with debris amongst the have been caught mainly in flight interception traps.
setae. Recorded from the Chatham Is, Dunedin district, and
Stewart I. [12] Staphylinidae: Omaiinae
Fig. 124 Omaliomimus albipennis (Kiesenwetter)
[8] Ptiliidae: Ptiliinae A medium-sized rove beetle 3.5 mm long, moderately
Fig. 118 Notoptenidium lawsoni Matthews convex and glossy, with head black, pronotum reddish-
An elongate-oval, convex feather-winged beetle 1.0-1.1 brown to black, elytra entirely yellow or black, and antennae
mm long, black with yellow legs and antennae. Prothorax with segments 1-5 yellowish brown, remainder darker.
with sides strongly curved; pronotum irregularly punctured; Occurs under decaying seaweeds on the shore. Recorded
Occurs under decaying seaweeds on the shore. Recorded from the Auckland Is, Campbell I., and (extralimital) Mac-
from the Auckland Is, Campbell I., and (extralimital) Mac- quarie l. [Redrawn after Steel 1964.]
quarie I.

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[12] Staphylinidae: Proteininae [12] Staphylinidae: Scaphidlinae
Fig. 125 Silphotelus nitidus Broun Fig. 131 Baeocera scutellaris (Redtenbacher)
A small beetle 1.3-1.6 mm long with body pale to dark A strongly convex, highly glossy beetle 2.4-2.5 mm long,
reddish-brown, and sometimes with yellowish humeri and with elytra long, truncate apically, exposing small, pale red
posterior elytral angles. Head with a punctiform fovea on pygidium; antennae and legs also pale red. Found under
either side; elytra covering nearly all of abdomen. Found loose bark of dead trunks and branches, under stones, or in
amongst leaf mould. Recorded from the South I. [Redrawn moist leaf litter. Some scaphidiine larvae are associated
after Steel 1966.] with fungi. Recorded from the Poor Knights Islands to
Southland.
[12] Staphylinidae: Pselaphinae
Fig. 126 Sagola laminata Broun [12] Staphylinidae: Piestinae
An elongate, slender, somewhat depressed beetle 2.2-2.3 Fig. 132 Parasiagonum hudsoni (Cameron)
mm long, glossy red, with elytra, antennae, palps, legs, and A subparallel, flattened beetle 6.0-8.0 mm long, dark brown
pubescence yellowish. A distinct 'waist' between prothorax with reddish legs. Head transverse, with antennae approx-
and elytra; elytra leaving 5 segments of abdomen exposed, imately as long as body; pronotum with anterior angles
without striae but with characteristic postbasal foveae. A projecting; elytra grooved. Found under bark of karaka
distinctive lamina on the underside of the head, just behind (Corynocarpus laevigatus), and probably saprophagous.
the mentum. Occurs in leaf litter, especially of sedges. Recorded from the Wellington area. [Redrawn after Steel
Fairly common in the Auckland area. 1950a.]

[12] Staphylinldae: Phloeocharinae [12] Staphylinidae: Osoriinae

Fig. 127 Pseudophloeocharis australis (Fauvel) Fig. 133 Nototorchus ferrugineus (Broun)
A small beetle 1.8-2.0 mm long, yellowish brown with head An elongate, parallel-sided, reddish-brown beetle 3.4-3.5
brown, or entirely brown. Abdomen bearing meshed mm long. Abdomen cylindrical, with tergites and sternites
microsculpture. Occurs on both main islands, and may be fused laterally; paratergites absent. Found in leaf litter and
adventive in New Zealand since it is known also from humus, most frequently under taraire-dominated vegeta-
Australia and New Caledonia. tion in the north of the North Island. Recorded from North-
land to Nelson.'
[12] Staphylinidae: Tachyporinae
Fig. 128 Sepedophilus sp. [12] Staphylinidae: Oxytelinae
A small, brightly coloured beetle 1.8-2.1 mm long, teddish Fig. 134 Carpelimus sp.
brown with head and posterior part of elytra dark brown, A small, flattened, dark brown or black beetle 1.5-1.7 mm
remainder of elytra yellowish, and abdomen yellowish long; abdomen broadening apically except for conical
posteriorly and brownish basally, bearing long protruding apex. Carpelimus species are saprophages occurring in
setae. Sepedophilus species all have a pubescent pro- forest litter, but may be attracted to dung and carrion. They
notum. They are common in forest litler, rotting logs, and are often abundant, but little studied; three species are
fungi and occur on both main islands. There are numerous known to be adventive.
Sepedophilus species in New Zealand, several of them not
yet named. [12] Staphylinidae: Euaesthetinae
Fig. 135 Agnosthaetus vicinus (Broun)
[12] Staphylinidae: Habrocerinae A small, reddish beetle 2.8-3.0 mm long with a dark
Flg. 129 Habrocerus capillaricornis (Gravenhorst) transverse spot in apical thitd of abdomen; forebody glossy
A medium-sized, flattened beetle 2.7-3.0 mm long, brown and sparsely pubescent or naked; elytra extremely short;
with forebody strongly glossy and abdomen moderately abdomen moderately glossy, with dense whitish pubes-
glossy. Distinct by its thin antennae with long setae and cence. Like all New Zealand euaesthetines it is endemic,
long, protruding abdominal setae. An adventive species, flightless, and inhabits forest litter. Recorded from the
recorded from mid Canterbury. northern part of the South Island and the Waikato.

[12] Staphylinidae: Aleocharinae [12] Staphylinidae: Pseudopsinae

Fig. 130 Aleochara hammondi Klimaszewski & Crosby Fig. 136 Pseudopsis arrowi Bernhauer
A robust, narrowly oval beetle 5.0-7.5 mm long, strongly A small, flattened, reddish-brown beetle 2.8-3.3 mm long.
glossy, black with appendages and central part of elytra Head bearing a shallow depression laterad of middle;
reddish brown. Like all species of Aleochara it has a distinct pronotum with 5 costae; elytra each with 2 longitudinal
but minute pseudosegment terminating the maxillary and costae and 2 lateral longitudinal carinae; abdomen with
labial palps. Aleochara occurs on both main islands and the slightly clubbed setae. Known from leaf litter in Nothofagus
Kermadecs; this species is restricted to the South I. forest. Pseudopsis species occur mainly in forest litter and
moss, and appear to be predators. Recorded from the Bay
of Plenty and the Nelson area.

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[12] Staphylinidae: Paederinae Elytra each with 3 or 4 broad, longitudinal ridges; legs
Fig. 137 Medon zeelandicus (Redtenbacher) fossorial; antennal club segments chunky; mandibles trian-
A medium-sized, subparallel, reddish beetle 4.5-5.0 mm gular. The sole species of an endemic genus with close
Iong, often with head and base of abdomen darker. Ter- relatives in Australia, New Guinea, and Chile. Larvae live in
minal segment of maxillary palps minute. Recorded from soil and probably eat grass roots. Adults fly during the
Foxton, North I. Paederinae occur in damp habitats in evening in spring and summer. Recorded from the Moko-
forests or near bodies of water, and are predators that hinau Is in the north to Ben Lomond in the southern lakes.
employ pre-oral digestion.
[14] Trogidae
[12] Staphylinidae: Staphylininae Fig. 144 Trox scaber (Linnaeus)
Fig. 138 Cafius litoreus Broun A medium-sized, predominantly fuscous carcass beetle
A Iarge, glossy black beetle 11-13 mm long, with some 6.4-6.5 mm long, oval and convex, rugose. Antennae short
segments of antennae and tarsi reddish, and pubescence and stout; legs slender, not obviously modified for digging.
yellowish. Abdomen coarsely punctured; tibiae covered An adventive European species recorded mostly in the
with blunt spines. Unlike most Cafius species, head much northern part of the North I., sometimes at light.
larger in male than in female. Found under decaying sea-
weed, where both adults and the large, pale brown larvae [15] Scarabaeidae: Aphodiinae
are predators on littoral amphipods. Adults fly readily, Fig. 145 Acrossidius tasmaniae (Hope)
especially in hot sunshine, and have been collected from A stout, glossy beetle 10-11 mm long, reddish brown to
October to March. Recorded widely from Cape Reinga to black, with appendages usually paler. Elytra each with 5
Dunedin. imptessed striae; protibiae each with 3 large teeth, and
other tibiae each with 1 larger and 1 smaller spine;
[13] Lucanidae: Aesalinae antennae with a 3-segmented lamellate club. Accidentally
Fig. 139 Ceratognathus parrianus Westwood introduced from south-eastern Australia, where it is a
A large, brown stag beetle 14-20 mm long, with purplish- common pasture pest. Larvae live in tunnels in the soil,
black blotches on pronotum and elytra. Antennae not emerging to feed at night. They are unusual in feeding on
elbowed, with a 3-segmented setose club. Larvae are C- grass leaves rather than roots, often dragging leaves into
shaped grubs occurring in decaying wood of karaka and the tunnel for consumption. Adults fly to light on warm
other trees. Adults are sometimes found in the larval summer evenings. Recorded from the North l, especially in
habitat, and occur throughout the year. Widely distributed, areas with light volcanic soil, and the South I. south of Mt
and some-times flying to light. Grey, Canterbury.

[13] Lucanidae: Lucaninae [15] Scarabaeidae: Aphodiinae

Fig. 140 Paralissotes reticulatus (Westwood), male Fig. 146 Phycochus graniceps Broun
Alarge,dkbrown raely dishtagbetl12-0m A small, strongly convex beetle 3.0-4.0 mm long, glossy
long. Head and pronotum with dull yellowish-brown, reddish brown to black. Legs with broad, spinose tibiae and
appressed scales arising from coarse punctures; elytra short tarsi modified for digging; head granulate frontally,
reticulated, with several narrow, elevated, non-squamose with small eyes; elytra almost globose, weakly striated.
bands; mandibles each with a strong apical tooth, a small Found in seaweed on beaches. Recorded from both main
subapical dorsal and ventral tooth, and a broad bifurcate islands.
basal lobe. Adults and larvae are commonly found in
decaying wood on the ground, and adults may also be [15] Scarabaeidae: Scarabaeinae
found under bark of living trees, particularly Nothofagus. Fig. 147 Onthophagus granulatus Boheman
Recorded from both main islands. A medium-sized beetle 5-8 mm long, with head and
pronotum largely bronze-green and elytra pale fulvous
[13] Lucanidae: Lucaninae mottled with dark brown. Elytral intervals with a row of
Fig. 141, 142 Geodorcus auriculatus (Broun) prominent glossy granules, each with an erect bristle. An
A Iarge, glossy, sexually dimorphic stag beetle 20-25 mm adventive Australian species often associated with cow
Iong. Elytra with erect, yellowish-brown scales arising from and horse dung. Known in New Zealand since the 1870s,
dense, coarse, circular punctures; head widest behind having probably arrived in stable manure with stock
eyes. Males have the head expanded and mandibles imported by early settlers. Occurs on both main islands,
bifurcate at the tips, with a dorsal tooth near the middle. with most records from the South l.
Recorded from the Waikato and Coromandel areas.
[15] Scarabaeidae: Melolonthinae
[13] Lucanidae: Lampriminae Fig. 148 Odontria giveni Watt
Fig. 143 Dendroblax earlii White Alarge,par-shped, ubescntbe l 13-5m long;
A chocolate-brown, glossy, heavily punctate stag beetle head red-brown or piceous; pronotum and elytra reddish
17-28 mm long, with conspicuous fulvous hairs ventrally. mottled with dark btown; scutellum datk brown; ventral

— 171 —
surface nacreous on coxae and sterna. Antennae in both exposed. Occurs in coastal areas, on sandy beaches
sexes 3-lamellate. As in all chafers, the legs are adapted for above high water matk, where larvae may be found under
digging. Adults are found on leaves of shrubs at night, and logs. Recorded from the North Island and the Nelson area
sometimes fly to artificial light. Larvae of most species of of the South I.
Odontria occur in forest soil, and occasionally in soil of
native tussock grassland.One of a group of alpine forest [16] Scirtidae
forms ranging throughout the South I. and onto the ranges Fig. 153 Amplectopus pallicornis Broun
of the North I. A small, elongate-oval, moderately pubescent, reddish-
brown beetle 1.9-2.0 mm long, with antennae and legs
[15] Scarabaeidae: Melolonthinae yellowish. Ventral surface with oval depressions into which
Fig. 149 Costelytra zealandica White head, antennae, and legs can be folded; antennae with a
A moderately large beetle 9-10 mm long, brownish black to regular 6-segmented club, and with proximal segments
pale brown, with edge of clypeus and eyes darker; irregular in form; legs folding together like the blades of a
antennae and (usually) elytra testaceous; pronotum pale pocket knife. Adults have been beaten from foliage of the
reddish brown to brown-black, and scutellum often reddish; tree fern Dicksonia squarrosa at Pelorus Bridge near
ventral surface pale brown. Pygidium partly exposed, Nelson. Originally found on Mt Pirongia and since collected
smooth and glabrous except for marginal bristles. Com- at Waipoua Forest, Hunua Range, Mt Arowhana (Gis-
monly known as the grass grub, a name more properly borne), Dawson Falls (Mt Egmont), and Takaka (Nelson).
applied to the larvae, which live in soil feeding on the roots
of grasses. Mature larvae pupate in the soil, usually in late [16] Scirtidae
August to late October, and the beetles emerge in Nov- Fig. 154 Veronatus tricostellus White
ember. On warm, still nights they fly, and alight on the A moderately large, narrowly oval, slightly depressed
leaves of various food plants such as turnip, plum, rose, beetle 9.0-10 mm long, pale brown with antennae and legs
apple, strawberry, and Aralia spinosa. After mating, usually yellowish brown. Elytra with 6 curved longitudinal costae;
on the food plants, females lay their eggs in moist soil. upper surface with fine, decumbent pubescence; head
Larvae hatch in 16-21 days and pass through three instars usually strongly bent downwards; penultimate tarsal
lasting from November to the following September. En- segment bilobed. Collected by beating shrubs. Widely
demic to New Zealand, from North Cape to Bluff, but not distributed on both main islands.
common in Northland and Auckland, and not prevalent in
native tussock grasslands. Causes most damage in the [17] Eucinetidae
South I., especially in Otago and Southland. Fig. 155 Eucinetus stewarti (Broun)
A small, broadly oval, glossy black beetle 3.5-3.6 mm long.
[15] Scarabaeidae: Melolonthinae Head and pronotum sparsely punctate; elytra with cross-
Fig. 150 Prodontria lewisi Broun striations; antennae cylindrical, 11-segmented, the last
Alarge,paler d ish-brownbe tl 15- 6m long; segment black; metacoxae large, fused to mesosternum;
antennae 4-lamellate in male and 3-lamellate in female; legs slender, the hind legs largest, with small spines. Adults
clypeus and frons densely and coarsely punctate; pro- and larvae probably feed on slime moulds. Discovered at
notum strongly convex; elytra striate, strongly convex, Boatmans near Reefton, and since found at Stratford; likely
truncate behind, with shoulders bearing a posterolateral to prove widely distributed wherever slime moulds occur.
depression. Known only from rough pastureland in a re-
stricted area of Cromwell, Central Otago. [18] Clambidae
Fig. 156 Clambus domesticus Broun
[15] Scarabaeidae: Dynastinae A minute, broadly oval beetle 1.0 mm long, pale to dark
Fig. 151 Heteronychus arator (Fabricius) brown, strongly glossy, sparsely pubescent. Head very
A moderately large, stout, convex, glabrous beetle 11-13 broad, deflexed beneath pronotum; antennae 8-seg-
mm Iong, strongly glossy black. Middle and hind femora mented, with a 2-segmented setose club; legs slender, with
each with 2 strong spurs. Commonly known as the black tibiae Iacking spurs. An adventive Australian species, fully
beetle, this species was accidentally introduced from South winged, and commonly occurring in lawn clippings, garden
Africa. Adults sometimes fly to light at night. Larvae feed on prunings, compost, and litter in native forest.
the roots of grasses, and may cause substantial damage to
lawns and pastures. Widely distributed in warmer areas of [19] Buprestidae
the North I. Fig. 157 Nascioides enysi Sharp
A moderately large, somewhat depressed, bullet-shaped
[15] Scarabaeidae: Dynastinae beetle 7-8 mm long, metallic green with 2 oblique yellow
Fig. 152 Pericoptus truncatus Fabricius spots; males with frons reddish. Elytra pointed apically.
Alarge,stoubetl 21-30m long,darkbown, ften Commonly known as the beech buprestid, the fully winged
chocolate brown medially. Legs broad, adapted for digging; adults feed in summer on leaves of Nothofagus, and are a
pronotum in males bearing a blunt apical horn; pygidium potential pest of beech forests.

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[20] Byrrhidae: Byrrhinae [25] Ptilodactylidae: Anchytarsinae
Fig. 158 Liochoria huttoni Pascoe Fig. 163 Byrrocryptus urquharti Broun
A moderately small, broadly oval, convex beetle 3.6-3.7 A moderately large, closely pubescent beetle 6-7 mm long,
m long,shin gblackwithgre nishrefl ctions.Anten ae with body medium brown and legs and antennae yellowish.
inserted on sides of frons, with a 3-segmented club; elytra Antennae serrate; scutellum heart-shaped; hind margin of
each with about 5 longitudinal rows of unconnected coarse pronotum with blunt teeth forming a transverse comb.
punctures; abdominal sternites behind metacoxae with a Adults have been collected between October and February.
large oval depression for reception of retracted hind legs. Larvae live in sand near the water's edge in river beds.
Adults are often found under stones surrounded by moss, Recorded from Auckland to mid Canterbury.
usually associated with larvae, and have been observed
feeding en alpine daisies (Celmisia). Recorded in the [26] Chelonariidae
southern South I. from the Grampian Mountains (Mac- Fig. 164a,b Brounia thoracica Sharp
kenzie Basin) to the great block mountains of Central A moderately Iarge, narrowly oval, slightly glossy beetle 5-
Otago, at altitudes from 350 m to over 3000 m. 6 mm long, black with tarsi yellowish-red and elytra violet or
purple-tinged; moderately densely punctate and pubes-
[21] Dryopidae cent. Head partially concealed from above; antennae with
Fig. 159 Parnida agrestis Broun segments 6-10 each bearing a long, inwardly produced
A moderately small, moderately glossy black beetle 3.9- lobe; pronotum hump-shaped in lateral view. The only
4.0 mm Iong, covered with irregular yellow hairs. Antennae known specimens are the original two collected by Broun
short, with a characteristic asymmetrical club; pronotum (locality unspecified) and another Malaise trapped at East
with a deeply impressed groove do either side near lateral Cape.
margin. One of a number of terrestrial Dryopidae (the
dryopids in other zoogeographical regions are mostly [27] Eucnemidae: Melasinae
aquatic). Adults and larvae occur commonly together in leaf Fig. 165 Neocharis simplex Sharp
litter of Nothofagus forests. A moderately small, elongate oval beetle 3.8-4.0 mm long,
black with greyish pubescence. Antennae long, the 2nd and
[22] Elmidae: Larainae 3rd segments smallest; elytra without definite striae. Found
Fig. 160 Hydora picea Broun on forest growth and in leaf litter and wood mould, and has
A moderately small, broad, slightly depressed, brownish- been taken in Malaise traps. Recorded from Kaihu in
black beetle 3.4-3.5 mm long. Head narrow, with prominent Northland to Hump Ridge in Southland.
eyes; antennae without a distinct club; posterior edges of
pronotum sharp; elytra with rows of punctures; body [28] Elateridae: Agrypninae
clothed with fairly short pubescence. Widely distributed in Fig. 166 Thoramus wakefieldi Sharp
riverbeds, on stones at the water's edge. A large, narrowly oval, blackish-brown click beetle 15-30
mm long. Sides of pronotum evenly curved anteriorly to
[23]
Limnichidae: Limnichinae blunt posterior angles; elytra each with about 5 striae;
Fig. 161 Limnichus nigripes Broun antennae slender; legs stout; metacoxae excavated
A small, oval, convex, glossy black beetle 2.3-2.5 mm long, posteriorly to receive retracted femur. One of our Iargest
clothed with yellowish scale-like setae arranged in wavy click beetles, quite variable in shape and size. Larvae ate
patterns, the setae finer and hair-like on pronotum. Anten- found in dead wood in forests, where they prey on larvae of
nae with a small 2-segmented club, inserted on base of other beetles such as cerambycids. Adults of this widely
frons. Occurs under stones in streams and at the margin of distributed species fly readily to light.
rivers. Also found in terrestrial habitats in moist litter and
under stones, but primarily an aquatic species. Recorded [28] Elateridae: Dentlcollinae
at Lowburn, just north of Cromwell, in the Motu R., Bay of Fig. 167 Amychus candezei Pascoe
Plenty, and in the Kereu Stm, East Cape. A Iarge, brownish click beetle 16-23 mm long, variegated
and variable in colour, resembling bark. Pronotum broader
[24] Heteroceridae: Heterocerinae than long, rounded at sides, and with prominent posterior
Fig. 162 Heterocerus novaeselandiae Charpentier angles; elytra narrowet than pronotum, tapeting posteriorly;
A moderately small, elongate, convex, brownish beetle body surface moderately rough. Occurs in crevices in rocks
3.9-4.0 mm Iong, bearing numerous bristles. Antennae and under logs and stones. Recorded from the islands of
short and thick; elytra with oval, yellowish-brown spots; the Cook Strait and the Chathams, this endemic New
tibiae flattened and spinose, blackish-brown. The only Zealand species is considered to be endangered.
species of heterocerid known from New Zealand, from a
few widely scattered records. [29] Lycidae
Fig. 168 Porrostoma rufipenne (Fabricius)
A moderately large, soft-bodied, elongate, depressed
beetle 9-10 mm long, black with elytra orange (often bright

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red in living specimens). Antennae deeply serrate; elytra males with a club consisting of 3 long branches, and in
with indefinite longitudinal ridges and large punctures female simple, with club segments smaller and slightly
between them. Recorded from the North I., especially asymmetrical. Described on the basis of a male and a
northern areas, and around Nelson in the South I. female found near Whangarei Heads and at Tairua.

[30] Cantharidae: Dysmorphocerinae [35] Bostrichidae: Dinoderinae

Fig. 169 Asilis fulvithorax Broun Fig. 175 Dinoderus minutus (Fabricius)
A medlum-sized, soft-bodied, fairly flat soldier beetle 4.8- A small beetle 2.0-3.0 mm long, dark brown with red elytra.
5.0 mm long, black with pronotum bright orange. Antennae Head with frons densely punctate; pronotum coarsely
slender and slightly serrate, inserted on frons just before asperate anteriorly, densely punctate posteriorly; elytra
eyes. Adults may be beaten from foliage of various trees irregularly punctate with short, erect setae. A pantropical
and shrubs, especially when in flower. Common in the pest species developing in bamboo, and sometimes in
North I. north of Waitomo, including the Hen and Chickens maize. Recorded from Auckland.
Is and Little Barrier.
[35] Bostrichidae: Lyctinae
[31] Derodontidae: Laricobiinae Fig. 176 Lyctus brunneus Stephens
Fig. 170 Nothoderodontus gourlayi Crowson A small to moderately large, brown beetle 2.0-7.0 mm long.
A small, broadly oval beetle 1.9-2.0 mm long, somewhat Upper surface with brown decumbent pubescence; head
depressed, brownish to black. Head with characteristic and pronotum with coarse, shallow punctures; elytra
depressions and with 2 ocelli, each next to margin of eye; striate, with fine punctation; pronotum slightly depressed
eyes prominent, coarsely faceted; antennae with a 3- medially; posterior margin of 4th ventrite with fine pubes-
segmented club; elytra with deep striae. Adults have been cence in both sexes. A cosmopolitan pest in a wide variety
collected from January to March. Discovered at Arthurs of timbers. Recorded from both main islands.
Pass, and reported from several localities between the
Nelson area and Capleston, near Reefton. [36] Anobiidae: Anobiinae
Fig. 177 Hadrobregmus magnus (Dumbleton)
[32] Jacobsonildae A medium-sized, reddish-brown beetle 7.0-8.0 mm long.
Fig. 171 Saphophagus minutus Sharp Head concealed by prothorax; antennae with a 3-
A small, elongate-oval, depressed, reddish-brown beetle segmented club consisting of elongate and asymmetrical
1.9-2.0 mm long. Pronotum approximately heart-shaped, segments; prothorax constricted posteriorly near base;
with a pronounced median depression and with coarse elytra with rows of punctures; most of body surface with
punctures; head less coarsely punctate; antennae straight, short pubescence. Adults and larvae occur in old, partly
with a loose 3-segmented club; elytra without distinct decayed rlmu logs and in rimu flooring and other timbers;
striae, but with punctures arranged in somewhat irregular, they usually bore into wet timber. Widely distributed, from
curved, longitudinal rows; legs tather short and stout. Northland to the Taieri Plains, Dunedin.
Collected in the Nelson area, at Picton, and near Mt Cook.
[36] Anobiidae: Ptininae
[33] Nosodendridae Fig. 178 Ptinus speciosus Broun
Fig. 172 Nosodendron zealandicum Sharp A small, broadly oval, spider-like beetle 2.3-2.4 mm long,
A moderately small, oval, strongly convex, glossy black with a deep brown and white stripe across elytra and
beetle 4.3-4.5 mm long, finely and closely punctate. Elytra additional white markings behind shoulders; body clothed
with rows of large punctures. Widely distributed in the North with long yellow hairs. Head usually strongly deflexed and
I., and also recorded from the South I. partially concealed from above; antennae without a club;
legs and antennae long and slender; elytra much broadet
[34] Dermestidae: Megatominae than prothorax. Adults occur in leaf litter. Widely distributed
Fig. 173 Trogoderma maestum Broun in the North I. and northern South I.
A small, broadly oval, slightly depressed beetle 2.9-3.0 mm
Iong, black, covered with black and whitish recumbent hairs [37] Trogossitidae: Rentoniinae
forming a characteristic pattern. Antennae of male bluntly Fig. 179 Rentonidium costiventris Crowson
serrate, those of female weakly clubbed. Adults may be A minute, oval, strongly convex, pale reddish-brown beetle
found on flowers, and larvae on feathers and dead birds. 1.3 mm long. Antennae and legs slender; head broad;
Recorded from the north of the North I., as far south as upper surface with fine pubescence; elytra without striae; 1
Okauia near Matamata. visible sternite with a median keel. Discovered in male
flowers of "Pinus insignis" (probably in error for P. radiata) in
[35] Bostrichidae: Euderinae Waipoua State Forest, Northland. Also collected in other
Fig. 174 Euderia squamosa Broun North I. localities in forest leaf litter, and occurring as far
A moderately small, elongate beetle 4.0-5.0 mm long, south as Nelson.
brown, covered with white and brown scales. Antennae in

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[37] Trogossitidae: Lophocaterinae coloured, mostly dull black with elytral shoulders or entire
Fig. 180 Grynoma regularis Sharp elytra pale yellowish, or elytra appearing yellowish with
A medium-sized, moderately convex, ventrally flat beetle various combinations of brown or black spots. Occurs on
5.3-5.4 mm long, elongate oval with sides mostly parallel. sandy beaches. Recorded from both main islands, the
Colour predominantly black; elytra with white marks Chathams, and the Kermadecs.
variegated with pale red, coarsely punctate but not striate;
Iong, whitish hairs covering most of dorsal surface; [41] Melyridae: Dasytinae
maxillary palps prominent, the last segment hatchet- Fig. 186 `Dasytes' subcyaneus Broun
shaped. Collected by beating forest foliage. Larvae are A medium-sized, soft-bodied, elongate beetle 3.8-4.0 mm
sometimes found under loose bark, where they probably long, slightly depressed, metallic blue. Head without a
feed on other insects; adults too are probably predacious. distinct neck; eyes prominent; antennae filiform, without a
Widely distributed. club; elytra pubescent, without striae. Common on flower-
ing shrubs, and when alerted takes to flight rapidly. Larvae
[37] Trogossitidae: Trogossitinae live under loose bark of dead branches. Occurs on both
Fig. 181 Lepidopteryx nigrosparsa (White) main islands, but with most records from the South I.
A large, elongate, parallel-sided, slightly convex beetle 11-
12 mm long, glossy greenish black. Head and pronotum [42] Nitidulidae: Nitidulinae
with depressed areas bearing yellowish-white scales; Fig. 187 Platipidia asperella Broun
elytra with 14 longitudinal keels. Adults have been found A medium-sized, broadly oval, slightly depressed beetle
undet bark and in the sheath of a fallen nikau palm leaf. 4.8-5.2 mm long, mostly dark brown, with pale irregular
Collected from September to April. Widely distributed, with markings. Eyes strongly convex; head, pronotum, and
records ranging from Little Barrier I. to Silverstream near elytra finely punctate; elytra without striae; male with hind
Wellington. femora expanded towards apex. Adults are found on tree
trunks, especially with flowing sap, and sometimes in the
[38] Chaetosomatidae base of fallen dead nikau palm leaves. Occurs en both main
Fig. 182 Chaetosoma scaritides Westwood islands, but with most records from the South I.
A Iarge, elongate, subparallel-sided beetle 13-14 mm long,
coarsely punctate, black with pale red shoulders and elytral [43] Monotomidae: Monotominae
apex. Dorsal surface with long, erect hairs; antennae and Fig. 188 Lenax mirandus Sharp
legs slender; pronotum subquadrate, with a strongly glossy A medium-sized, narrowly elongate, subparallel beetle 4.0-
median elevation. Collected from under bark of logs and 6.0 mm long, usually dark brown to nearly black, with
standing dead trees of various species. Widely distributed. appendages paler, usually rust-brown; glossy, with minute,
sparse and inconspicuous pubescence. Head short, as
[39] Cleridae: Thanoclerinae broad as thorax, with 2 deep cavities behind each eye;
Fig. 183 Metaxina ornata Broun labrum concealed; pronotum narrowly elongate, with front
A medium-sized, slightly flattened beetle 4.0-6.0 mm long, angles sharp and slightly produced, lateral carinae sharp,
brown with whitish-yellow, sinuate, irregularly shape elytral and with 2 rows of Iarge, deep punctures in medial part of
spots. Dorsal surface with long, erect sparse setae; head disc; elytra narrowly elongate, with 8 punctate grooves.
and pronotum with longitudinally elongate punctures, and Collected under bark in beech (Nothofagus) forests, and
elytra with round ones. Originally described from Broken sometimes by beating forest growth. Widely distributed,
River, Canterbury. We have seen additional speclmens especially in the South I.
from Dun Mountain, near Nelson, and an undetermined
specimen of the same genus from Northland. [43] Monotomidae: Monotominae
Fig. 189 Monotoma spinicollis Aubé
[39] Cleridae: Enopliinae A small beetle 2.1-2.2 mm long, entirely rust-brown, or dark
Fig. 184 Phymatophaea violacea (Fabricius) brown with rust-brown edges and appendages. Head and
A medium-sized, elongate-oval, slightly depressed beetle pronotum with coarse, broad, dense punctation; head
6.5-7.0 mm long, mainly violet, with distinct yellow mark- abruptly constricted behind, with protruding eyes; pro-
ings on elytra. Eyes prominent; antennae with an elongate notum with anterior angles produced and lateral margin
and asymmetrical 3-segmented club; 2 tarsal segments serrate; elytra shortened, with rows of punctures; pubes-
strongly lobed; elytra coarsely punctate, without striae, cence short, inconspicuous, yellowish. Adults of this intro-
clothed with long, protruding hairs. Adults occur in summer duced European species are common in lawn clippings,
on flowering trees and shrubs, where they probably prey on garden litter, and compost, in paddocks where sheep
other insects. shelter, and in chicken straw. Recorded from Auckland.

[40] Phycosecidae [44] Phloeostichidae: Agapythinae

Fig. 185 Phycosecis limbata (Fabriclus) Fig. 190 Agapytho foveicollis Broun
A small, elongate-oval beetle 2.5-3.0 mm long, variably A moderately small, narrowly elongate beetle 3.0-3.2 mm

—175—
long, rust brown with dark brown spots and 2 pale, [47] Laemophloeidae
transverse spots on elytra. Moderately glossy, sparsely Fig. 195 Microbrontes lineatus (Broun)
pubescent, with depressions on pronotum and elytra and A small, flattened beetle 2.8-3.0 mm Iong, brown with a
with inconspicuous tubercles; antennae bead-like, with a reddish tinge, slightly glossy, with inconspicuous short
weakly defined 3-segmented club. Adults have been pubescence. Head elongate, wifh a median ridge and 2
collected from December to February. Originally taken at lateral ridges; antennae with scape enlarged and asym-
Routeburn, but also known from the Hollyford Vly, Balloon metrical, pedicel broadened apically, and remaining seg-
Hut (Mt Arthur Tableland, Nelson), Takaka Hill, Mt Robert ments strongly elongate; pronotum broadest apically, with
(Nelson Lakes N.P.), upper Wairoa Vly, and Makarora. 2 sublaferal ridges; elytra with 10 longitudinal ridges.
Known from fhe North I. and Three Kings Is.
[44] Phloeostichidae: Priasilphinae
Fig. 191 Priasilpha obscura Broun [48] Phalacridae: Cyclaxyrinae
A medium-sized, broadly oval, flattened beetle 4.7-5.0 mm Fig. 196 Cyclaxyra i mpressa Broun
Iong, narrowed posteriorly, blackish brown, coarsely A small, strongly convex beetle 2.0-2.2 mm Iong, glossy
punctate. Upper surface with greyish pubescence grouped black with legs and antennae reddish chestnut. Head and
in clumps on elytra; elytra without striae, slightly explanate; elytra sparsely punctate; antennae with a 3-segmented
antennae with a distinct 3-segmented club; antennae and club; elytra not striate. Presumably fungivorous, occurring
legs slender. Adults and larvae occur in leaf litter of beech in sooty moulds on Nothofagus and Olearia. Known from
(Nothofagus) forests. Widely distributed on both main both main islands.
islands.
[49] Cavognathidae
[45] Silvanidae: Brontinae Fig. 197 Zeonidicola chathamensis Watt
Fig. 192 Brontopriscus pleuralis Broun A small, elongate-oval, moderately depressed beetle 2.7-
A moderately Iarge, narrowly oval, flattened, brown beetle 3.2 mm Iong, dark brown with legs and antennae reddish
7.5-8.0 mm Iong. Head slightly narrower than pronotum; brown; surface relatively dull, with fine microsculpture.
antennae with scape extremely elongate, as Iong as 3 or 4 Head densely punctate; antennae with a loose 3-seg-
following segments combined; pronotum with deep inden- mented club; pronotum with sides weakly angulate just
tations on lateral margins; elytra oval and flat laterally, with behind middle; elytra elongate, widest at mid-length; wings
longitudinal rows of punctures; dorsal surface sometimes vestigial; scutellum transverse; legs stout, with protarsi and
bearing irregular pieces of plant material adhering to sappy mesotarsi (segments 1-4) expanded in male. Occurs in
secretion. Adults and larvae occur under bark of dead tree birds' nests on the Chatham Is.
trunks and branches of native trees. Common in most parts
of the North I., with a few records from the South I. [50] Cryptophagidae: Cryptophaginae
Fig. 198 Thortus ovalis Broun
[45]
Silvanidae: Silvaninae A small, oval, convex beetle 1.7-1.8 mm Iong, reddish
Fig. 193 Oryzaephilus surinamensis (Linnaeus) brown with shoulders of elytra and patches near elytral
A small, narrowly elongate beetle brown 2.8-3.2 mm Iong, apex paler; most of upper surface with slender setae.
often with rust-brown appendages; dorsal surface covered Antennae with a loose 3-segmented club, fheir inserfions
with short, yellow, recumbent hairs. Head elongate, with depressed into an elevated surface and separated by a
temples as long as diameter of eye; antennae with a 3- longitudinal carina; lateral carinae of prothorax without
segmented club; pronotum with 6 teeth on either side; serration; elytra with fine punctation and without striae;
elytra with longitudinal rows of punctures. A cosmopolitan wings absent. Occurs in leaf litter, especially in Nothofagus
pest of stored products, adventive in New Zealand. Adults forests. Discovered at Boatmans near Reefton, and later
and larvae infest cereals, dried fruits, and oilseeds. Known found on Stephens I. and more widely on the western side
from both main islands. of the South I.

[46] Cucujidae [51] Languriidae: Xenoscelinae

Fig. 194 Platisus sp. Fig. 199 Hapalips prolixus (Sharp)
Alarge,flatbe l 15-20m long,darkbown, ith ead A medium-sized, narrowly elongate beetle 5.0-6.0 mm
and pronotum nearly black, and elytra uniformly dark brown Iong, subparallel to slightly convex, yellowish-brown.
or reddish apically, coarsely punctate. Head transverse, Antennae with an elongate 3-segmented club; pronotum
posteriorly slightly broader than pronotum; mandibles withouf a basal impression or groove; elytra with rows of
strong, curved, with inner teeth; antennae moniliform, wifh punctures. Adults and larvae occur on dead leaf tissue of
basal and 3rd segments strongly elongate and 2nd tree-ferns and nikau palm. Known from the North I. and the
segment reduced and beadshaped; pronotum with serrate northern part of the South I.
lateral margins. An endemic New Zealand species known
only from the Three Kings Is.

-176-
[52] Erotylidae: Dacninae [55] Endomychidae: Holoparamecinae
Fig. 200 Thallis polita White Fig. 205 Holoparamecus tenuis Reitter
A medium-sized, elongate-oval, convex beetle 5.8-6.0 mm A small, elongate-oval, reddish-brown beetle 1.5-1.7 mm
long, blackish-brown, strongly glossy. Head with prominent Iong. Antennae with a Iarge, 2-segmented club; pronotum
eyes and two deep grooves; antennae with a loose 3- convex, restricted behind to a distinct 'waist', wlth a series
segmented club; elytra with irregular longifudinal rows of of laterally elongate foveae forming a basal impression;
fine punctures. Adults are found on bracket fungi and elytra pubescent, without striae; legs slender, with tibial
occasionally in leaf litter or under bark. Widely distributed in spurs. Associated with leaf litter under rotting logs and with
the North I. fungi. Fairly common, and probably widely distributed.

[53] Bothriderldae: Anommatinae [56] Coccinellidae: Coccinellinae

Fig. 201 Anommatus duodecimstriatus ( Müller) Fig. 206 Coccinella leonina Fabricius
A small, subparallel beetle 2.0-2.2 mm long, dorsally A medium-sized, hemispherical ladybird 5.3-5.5 mm long,
glossy, depigmented, pale brown, somewhat flattened, with with many conspicuous, regular orange markings on a
short, sparse and inconspicuous setae. Head partially black background. Adults and larvae feed mostly on aphids
concealed by pronotum, distinctly narrower than apical occurring on grasses and tussock, but are known from a
pronotal margin; eyes reduced; antennae 10-segmented, wider range of plants. Recorded widely over both main
with a Iarge 1-segmented club; pronotum trapezoldal, with islands and from smaller islands close offshore, from near
large, longitudinally elongate punctation; elytra with narrow sea level to subalpine.
epipleura, and with 6 rows of setose punctures on either
side. Adults recorded from rotten wood buried in earth, [57] Corylophidae: Sericoderinae
under deeply buried stones, and from deep pitfall traps Fig. 207 Anisomeristes sharpi Matthews
sealed off at the top. An introduced hypogeal European A small, densely pubescent, glossy, bright orange beetle
species, recorded from Lynfield, Auckland. 1.2-1.4 mm Iong. Head concealed by front of pronotum;
antennae 10-segmented, with a 3-segmented club; pro-
[53] Bothrideridae: Bothriderinae notum with hind angles strongly produced posteriorly;
Fig. 202 Ascetoderes obsoletus (Broun) elytra without striae, slightly truncate, exposing part of
A medium-sized, elongate-oval, slightly depressed beetle pygidium; 2nd tarsal segment bilobed. Found on and under
5.0-6.0 mm long, dark brown to black with most of surface leaf litter at the base of sedges and flax, and in coastal cliff
coarsely punctate. Antennae inserted in front of eyes, with vegetation around Auckland.
a 2-segmented club; pronotum with a medial impression;
elytra striate; legs stout, the segments simple. Adults are [58] Corticariidae: Lathridiinae
occasionally found under bark of dead tree trunks. Col- Flg. 208 Enicmus caviceps Broun
lected November to March. Recorded from the Nelson area A small, elongate-oval, glossy black beetle 1.9-2.0 mm
(several records) to Invercargill. Iong. Antennae straight, wifh a 3-segmented black club;
elytral intervals convex, with intervals 3 and 5 costate
[54] Cerylonidae: Euxestinae posteriorly, and interval 7 costate throughout. Occurs in
Flg. 203 Hypodacnella rubriceps (Reitter) leaf litter at the base of sedges. Found frequently in the
A small, oval, strongly convex, glabrous beetle 2.0-3.0 mm Auckland area, and apparently widely distributed.
long, dark brown and glossy with appendages reddish.
Head slightly inclined, narrower than pronotum; antennae [59] Mycetophagidae: Mycetophaginae
10-segmented with a 2-segmented round club, and pedicel Fig. 209 Triphyllus hispidellus ( Broun)
and scape asymmettical; pronotum trapezoidal, sinuate A small, oval, slightly depressed beetle 1.8-1.9 mm long,
basally; elytra with fine punctures; abdomen with 1st black with antennae and legs yellowish; fine, short black
ventrite bearing femoral lines; 5th ventrite with fine lateral pubescence and paler-coloured spots on elytra. Antennae
expansions. Adults occur in leaf litter and other organic slender, with a 3-segmented club, the segments simple.
litter. Predominantly from the North I., but some records Beaten or swept from manuka (Leptospermum scoparium)
from the Nelson area. and other shrubs; also on rushes. Adults were originally
discovered at Whangarei Heads, and later reported from
[55] Endomychidae: Mycetaeinae The Noises Is, Mokohinau I., and several localities in the
Fig. 204 Mycetaea subterranea (Fabricius) Nelson area.
A small, oval, hairy, reddish beetle 1.7-2.0 mm Iong, with
sparse protruding hairs. Pronotum sinuate basally, with an [60] Archeocrypticidae
impressed lateral line on either side; elytra elongate, with Flg. 210 Archeocrypticus topali Kaszab
distinct shoulders. An adventive species, known as a minor A small, uniformly oval, finely pubescent beetle 2.7-3.5 mm
pest in temperate regions, feeding on moulds in stores and Iong, dark brown to nearly black. Antennae with a 3-
cellars. Recorded from the Nelson area. segmented club; prosternal process abruptly expanded
apically. Introduced into New Zealand from South America.

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[61] Ciidae: Ciinae tentatively assigned to this species. Recorded from both
Fig. 211 Cis zeelandicus Reitter main islands.
A small, subcylindrical beetle 1.8-2.0 mm long, dark brown,
with short bristles. Anterior margin of head with 2 blunt [65] Colydiidae: Colydiinae
teeth; antennae with a 3-segmented, loosely formed club; Fig. 216 Rhizonium antiquum Sharp
pronotum broadly rounded anteriorly, approximately A small, narrowly oval beetle 1.9-2.0 mm long, slightly
straight at base; elytra with confused rows of punctures. depressed, with upper surface bearing dense and coarse
Adults and larvae are often found together in cylindrical punctures. Antennae with a loose 3-segmented club; basal
tunnels in artists' shelf fungus (Ganoderma applanatum), antennal segment fully visible from above; elytra mostly
but adults also occur in the forest canopy and less subparallel; legs short. Adults occur in rachides of dead
frequently in litter. A fairly common species, widely dis- leaves of the tree fern Cyathea dealbata, and in decaying
tributed. wood of broadleaved woody plants. We have seen
specimens from the North I. only.
[62] Melandryidae: Melandryinae
Fig. 212 Hylobia nubeculosa Broun [66] Ulodidae
A medium-sized, elongate-oval beetle 4.3-4.4 mm long, Fig. 217 Syrphetodes marginatus Pascoe
orange-brown variegated with dark brown, with shorter and A moderately Iarge, broadly oval beetle 8.0-9.0 mm long,
longer pubescence. Head bent under pronotum; antennae slightly depressed, with integument brown, but variable in
without a distinct club; hind legs stouter than the others, shape and colour; upper surface with patches of scales
bearing Iong, minutely serrate tibial spurs; metacoxae large forming a variable pattern. Head partially concealed by
and flat. Adults may be found adhering to the underside of pronotum; eyes almost circular; antennal insertions ex-
decaying logs, where larvae probably feed on fungi. posed; antennae slender, without a club; pronotum with
Recorded from both main islands, from near sea level to front angles strongly produced anteriorly; elytra broad,
montane habitats. much broadet than pronotum. Adults may be found clinging
to the underside of logs, but are difficult to spot because of
[63] Mordellidae: Mordellinae their cryptic colour pattern. Larvae live in rotten logs.
Fig. 213 Mordella antarctica White Recorded from the Coromandel to Mt Hope near Nelson.
A large pintail beetle 13-17 mm long, black with charac-
teristic white markings on elytra. Antenna slightly serrate; [67] Chalcodryidae
pronotum sinuate basally; elytra rounded posteriorly; Iast Fig. 218 Chalcodrya variegata Redtenbacher
abdominal segment prolonged into a terminal spine. Adults Alarge, osybetl15-6m long,paletodrkgenor
occur frequently on flowering manuka (Leptospermum greenish brown, often with metallic reflections, and brown
scoparium) or rata (Metrosideros robusta) in full sun, are spots on elytra, but colour pattern variable; underside dark
very active and good jumpers, and take to flight readily greenish or brown; apex of femora, tibiae, and tarsi
when disturbed. Widely distributed. darkened; antennae pale brown; dorsal surface of head
parallel to margins of eyes, pronotal sides, and disc of
[64] Rhipiphoridae: Pelecotominae elytra with small patches of coarse yellowish pubescence.
Fig. 214 Rhipistena lugubris Sharp Antennae slender, lacking a club, with insertions exposed;
A medium-sized, elongate-oval, blackish-brown antlered vertex between eyes bearing an impunctate area; elytra
beetle 6-8 mm long. Antennae lamellate in male; eyes subparallel for most of length; legs slender. Confined to
kidney-shaped, partly surrounding base of antennae; high-rainfall forests, mostly Nothofagus. Recorded in the
pronotum with sides arcuate and base sinuate; elytra with North I. from the central volcanic plateau and around Wel-
rounded apices barely covering wings, non-striate, with lington, and in the South I. from Westland and Fiordland.
definite longitudinal ridges. Adults have been collected by
beating vegetation in summer; they fly very readily. Larvae [68] Tenebrionidae: Lagriinae
apparently are predatory, in dead wood. Recorded from the Fig. 219 Chaerodes trachyscelides White
North I. and the northern South l. A medium-sized, robust, strongly convex beetle 6.5-8.6
mm long; colour pattern very variable, from unicolorous
[65] Colydiidae: Colydiinae dark brown to tan with darker markings on pronotum and
Fig. 215 Pristoderus antarcticus White elytra. Protibiae strongly expanded anterolaterally, with
A moderately Iarge, broad, rather convex beetle 7.8-8.0 tarsi subterminal; meso- and metatibiae with pairs of stout,
mm long, with most of surface bearing short bristles. blunt, peglike apical spines; metatarsi angled upwards and
Antennae with a 3-segmented club; lateral margins of backwards — all modifications for moving on loose sand.
pronotum with 4 prominent teeth; elytra mostly subparallel, Recorded from sandy coasts from the Far North to Stewart
with fairly irregular, longitudinally arranged elevations; I., often under seaweed or other beach wrack.
abdominal sternites fully movable. Adults occur on stand-
ing dead tree trunks and sometimes under loose bark, often
with moulds. Larvae found in association with adults are

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[68] Tenebrionidae: Pimeliinae quadrate; elytra broad-shouldered, coarsely punctate,
Fig. 220 Actizeta albata Pascoe without strlae. Found on forest growth in summer. Adults
A small, pale-coloured darkling beetle 2.8-3.0 mm long; are pollen feeders, and larvae probably live under loose
upper surface covered with pale greyish-white scales. bark of dead branches. Widely distributed from Auckland to
Head partly concealed by pronotum; eyes coarsely fac- Southland.
eted; pronotum broad, with front angles broad and slightly
produced anteriorly; elytra with striae; forelegs broadly [71] Pyrochroidae: Pilipalpinae
expanded for digging, and hind legs bearing long bristles, Fig. 225 Exocalopus pectinatus Broun
an adaptation for walking on sand. Lives in loose sand on A medium-sized, elongate beetle 4.5-5.5 mm long, sub-
beaches throughout New Zealand. parallel-sided with elytra slightly broadening posteriorly,
glossy black, with fine pubescence. Head narrowed behind
[68] Tenebrionidae: Tenebrioninae eyes, forming a moderately narrow neck; eyes prominent;
Fig. 221 Mimopeus elongatus (Brême) maxillary palp with Iast segment broadly triangular; anten-
Alarge,longate-ovl,.moderatlyconvexdarklingbetl nae serrate in female, pectinate in male; pronotum slightly
11-15 mm long, black or reddish-black; upper surface transverse, with several impressions; elytra with prominent
closely punctate, the interstices between punctures shoulders, coarsely punctate, without striae. Occurs in
variable; surface dull or glossy, with or without apparent forested areas of Gisborne, Taupo, Taranaki, Wellington,
microsculpture. Head with expanded margin below each and Nelson to Southland, mainly at higher altitudes.
eye forming a canthus; antennae with an indistinct 5-
segmented club; pronotum transverse, the front angles [72] Salpingidae: Salpinginae
produced, without striae; legs slender. Adults usually feed Fig. 226 Salpingus bilunatus Pascoe
on dead vegetable debris, but in dry conditions feed on A small, elongate-oval beetle 2.0-2.2 mm long, with a
living plant tissue. Widely distributed, particularly in coastal distinct 'waist' between pronotum and elytra, yellowish
localities under creeping coastal plants such as pohuehue. brown with 3 black spots on elytra, the spots variable in
ln Auckland and on some offshore islands common under shape, and sometimes fused together. Antennae with a
stones in craters of extinct volcanoes. blackish 4-segmented club; pronotum with large, deep
punctures; elytra without distinct striae, but with slightly
[69] Prostomidae irregular rows of coarse punctures. Adults frequently found
Fig. 222 Dryocora howitti Pascoe by beating scrubby forest growth in summer. Widely
A medium-sized, elongate beetle 6.0-8.0 mm long, sub- distributed.
parallel, depressed, usually reddish brown. Head with
posteriorly produced lobes behind eyes; pronotum elon- [72] Salpingidae: Inopeplinae
gate; elytra faintly punctate-striate. Adults and larvae may Fig. 227 Diagrypnodes wakefieldi Waterhouse
be found in logs of rimu and hinau, in damp crevices be- A moderately large, elongate, strongly depressed beetle
tween wood blocks in the moist, deep red stage of decay. 5.0-10 mm long, glabrous, pale brown with darker elytra.
Widely distributed. Head elongate behind small eyes; antennae filiform, with
scape enlarged; pronotum strongly narrowed posteriorly,
[70] Oedemeridae: Oedemerinae truncate apically; elytra shortened, slightly convex, with
Fig. 223 Thelyphassa lineata (Fabricius) apex broadly rounded, leaving at least 4 abdominal seg-
Alarge, longate-oval xbetl 18-20m long,slihty ments exposed. Found with larvae, also very depressed,
depressed, with integument soft. Head produced in front of under loose bark of dead trunks and branches, feeding on
eyes; antennae long, pale yellowish brown, without a club, decayed cambial tissues. Widely distributed, and recorded
the segments subcylindrical, with scape fully exposed; from the Chatham Is.
pronotum with a dark median stripe; elytra with dark lateral
longitudinal stripes, coarsely and closely punctate, with 2 [73] Anthicidae: Anthicinae
indistinct longitudinal costae. Adults are found on flowers of Fig. 228 Anthicus hesperi King
native shrubs, where they probably feed on pollen and A medium-sized ant beetle 3.5-4.0 mm long, black with
nectar. Occasionally they occur in rotten wood on the forest orange-yellow elytral spots and posterior pronotum;
floor. Larvae live in wet rotten wood. Widely distributed, with pubescence sparse. Head subparallel behind eyes, then
records from Stewart I. and the Chathams. abruptly constricted into a nartow neck; eyes small;
antennae without a club; maxillary palps with basal 2
[71] Pyrochroidae: Pilipalpinae segments small and cylindrical; pronotum narrowed
Fig. 224 Techmessa concolor Bates posteriorly; elytra with pronounced shoulders, almost twice
A medium-sized, elongate, subparallel-sided beetle 5.0- pronotal width; metacoxae sepatated by more than one
6.0 mm long, slightly depressed, covered with fine pub- coxal width. Recorded in Auckland from a paddock,
escence. Head narrowed behind eyes, forming a broad heaped-up coarse prunings, and a stream bed in the bush.
neck; eyes prominent; antennae serrate; Iast segment of Also known from Nelson, and probably widely distributed.
maxillary palp broadly triangular; pronotum small, sub-

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[73] Anthicidae: Lagrioidinae mm long, slightly depressed, with body soft and fragile,
Fig. 229 Lagrioida brouni Pascoe pubescent, pale yellow with dark brown markings on elytra.
A medium-sized ant beetle 4.0-5.5 mm long, orange with Head slightly transverse, deflexed; eyes C-shaped, partly
reddish head, or head and pronotum brown with elytra surrounding antennal base; antennae filiform; maxillary
mottled yellow and brown; legs and antennae yellow to palps with last segment broadly expanded; pronotum
orange; antennal club sometimes darker. Associated with strongly transverse, with 2 small basal impressions; elytra
coastal sand dunes, and may be found beneath beach with confused punctation, Iacking striae; tibial spurs small,
debris. Recorded from both main islands, Stewart I., and with minute pubescence. Frequently found on foliage and
the Chathams. flowers in summer. Widely distributed on both main islands.

[73] Anthicidae: Lemodinae [76] Cerambycidae: Prioninae

Fig. 230 Cotes crispi (Broun) Flg. 234 Prionoplus reticularis Whlte
A moderately small ant beetle 2.9-3.1 mm long; head, The heaviest New Zealand beetle, 40-50 mm long, though
pronotum, and base of elytra reddish; remainder of elytra like most other wood-borers variable in size depending on
orange, with a brown transverse spot medially; appen- the nutrition of the larva, dark brown with a reticulate
dages yellow, glossy; pubescence moderately dense and yellowish pattern on elytra. Body elongate, somewhat
yellow. Head narrowly constricted behind eyes into a depressed; antennae almost as long as body; pronotum
narrow neck; eyes large, protruding; antennae incrassate with lateral margin bearing spines; ventral surface covered
apically but without a distinct club; basal 2 segments of with dense pubescence; legs long and slender. Larvae
maxillary palp with small projections; pronotum constricted attack mainly dead conifers, especially radiata pine and
near base; elytra without distinct shoulders, less than twice rimu, but are reported also from wattle; they require wood
pronotal width. Collected under Phormium tenax, Astelia with a high moisture content. Adults do not feed, utilising fat
banksii, and Gahnia setίfolίa. Originally described from reserves laid down by the larvae. Huhu grubs were once a
Parau, and widely recorded in the North I. significant food resource for Maori. Recorded from the
Kermadec Is to Stewart I., often at light.
[73] Anthicidae: Macratriinae
Fig. 231 Macratria exilis Pascoe [76] Cerambycidae: Cerambycinae
A medium-sized ant beetle 4.0-4.2 mm long, dark brown to Fig. 235 Zorion sp.
nearly black with basal antennal segments and legs A small longhorn 6.0-7.0 mm long, reddish brown with
reddish brown; pronotum and elytra coarsely punctate. yellowish spots on elytra, occurring on the Canterbury
Head abruptly constricted behind eyes into a narrow neck; Plains. There are several endemic Zorion species in New
eyes Iarge, protruding; antennae with a loosely formed, Zealand, all small and strikingly colourful, pubescent or
indistinct, 3-segmented club; maxillary palps with basal almost glabrous (e.g., blue with orange spots, orange with
segments bearing triangular projections; pronotum oval, yellow spots), but because of their small size they are
with base and apex broad; elytta with pronounced frequently overlooked. Adults can be found feeding in
shoulders, almost twice pronotal width; metacoxae almost native flowers. Recorded from both main islands.
touching. Beaten from foliage in midsummer, and found on
sedges. Recorded from near sea level around the North I. [76] Cerambycidae: Cerambycinae
Fig. 236 Oemona hirta (Fabricius)
[74] Aderidae A large,subcylindaohret15-2mlong,
Fig. 232 'Xyiophilus'nitidus Broun matt dark brown with scutellum and eyes orange. Antennae
A small, elongate-oval, slightly depressed beetle 1.9-2.0 slender, as long as body or nearly so; pronotum with
mm long, black with legs and antennae yellowish, punctate, irregular transverse grooves; elytra without striae, covered
clothed with greyish recumbent hairs. Head strongly with coarse brownish pubescence. Adults collected by
narrowed behind eyes into a distinct neck; eyes prominent, beating forest growth during summer, or occasionally at
coarsely faceted; antennae slightly broadening apically, light. Larvae attack many woody plants, boring into live
with terminal segment enlarged and oval; pronotum small, wood of citrus trees (hence the common name 'lemon tree
half of elytral width; elytra broad, with a depression behind borer'), grape, and various native plants, especially
shoulders, punctate but without striae; hind tibiae with rangiora. Recorded mostly from the northern half of the
sharp apical processes. Adults occur on foliage of Meryta North I. and around Nelson in the South I.
sinclairii (Three Kings puka) and other shrubs, and one
specimen was found in a robin's nest. Discovered at [76] Cerambycidae: Lamiinae
Northcote, Auckland, and subsequently recorded from Fig. 237 Xylotoles costipennis (Breuning)
Lynfield and from Lady Alice I., Hen and Chickens group. A moderately large, narrowly oval, convex longhorn beetle
8.0-14 mm long, with prothorax cylindrical, subglabrous,
[75] Scraptiidae pale brown to dark brown with patches of paler, yellowish
Fig. 233 Nothotelus usitatus (Broun) pubescence on sides of pronotum and 4 narrowly elongate
A moderately small, elongate-oval leaping beetle 2.9-3.0 spots on elytra. Head oriented downwards; eyes strongly

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emarginate; antennae longer than body, inserted in [77] Chrysomelidae: Cryptocephalinae
emargination of eyes; pronotum cylindrical; elytra with Fig. 242 Arnomus brouni Sharp
inconspicuous shoulders, broadest in basal half; legs stout. A moderately small, subcylindrical beetle 2.5-3.5 mm long,
Recorded mainly from Auckland, Coromandel, and North- rust brown with 6 apical antennal segments darker, and
land. often darker head, legs, and body margins; densely and
coarsely punctate. Eyes evenly convex; pronotum emar-
[77] Chrysomelidae: Bruchinae ginated and without teeth; pygidium usually partly exposed.
Fig. 238 Bruchidius villosus (Fabricius) Collected by beating manuka (Leptospermum scoparium)
A small, ovate bean beetle 1.7-3.5 mm long, pubescent, in flower. Recorded from around Auckland, and in the
slightly glossy. Head almost entirely concealed by pro- Nelson area.
notum; antennae with Iast 4 or 5 segments enlarged;
pronotum semicircular anteriorly, sinuate basally and with [77] Chrysomelidae: Eumolpinae
pubescence forming sinuate patterns in medial part of disc; Fig. 243 Eucolaspis brunnea (Fabricius)
elytra broad and shortened, exposing pygidium, with A medium-sized, elongate-oval beetle 4.8-5.0 mm long,
distinct striae; metatibia with an inconspicuous posterior yellowish brown but variable, ranging from dark brown to
spine. Introduced from the United Kingdom for control of brownish-ochreous, usually with a blackish stripe down the
broom (Cytisus scoparius). suture and one on either side of elytra. Upper surface
strongly punctate; antennae filiform. Common on manuka
[77] Chrysomelidae: Chrysomelinae (Leptospermum scoparium), especially when flowering,
Fig. 239 Chrysolina hyperici Forster and on the foliage of other native trees and shrubs. Bronze
A medium-sized, broadly oval leaf beetle 5.0-7.0 mm long, beetle larvae inhabit soil, especially under manuka. Widely
metallic green or blackish metallic green. Head deeply distributed.
inserted into prothorax; antennae short, broadly separated
basally; pronotum transverse, slightly emarginate apically, [78] Nemonychidae: Rhinorhynchinae
sinuate basally, almost equal in width to elytra; elytra broad, Fig. 244 Rhinorhynchus rufulus (Broun)
rounded posteriorly, with fine, dense punctation and large, A small weevil 2.0-2.8 mm long, narrowly oval, rust brown
sparse punctures in double confused rows; legs stout, with to dark brown with moderately dense, decumbent, yellow-
all tibiae similarly developed. Introduced into New Zealand ish-grey pubescence. Head abruptly constricted in front of
for biological control of St John's wort (Hypericum), and eyes into a narrow, apically flattened rostrum; antennae
now established. inserted in apical half of rostrum, straight, with a loose 3-
segmented club; prothotax as long as wide or longer in
[77] Chrysomelidae: Galerucinae (Alticini) male, as wide as long or slightly wider in female; elytra
Fig. 240 Agasicles hygrophila Selman & Vogt distinctly striate; hind wings well developed. Adults are
A medium-sized, elongate-oval flea beetle 4.5-5.0 mm associated with Podocarpacae, and larvae are pollen
long, black with base of antennae yellowish red, and with feeders in male inflorescences of podocarps. Recorded
longitudinal U-shaped whitish-yellow spots on elytra; upper from the Far North to Stewart I.
surface densely punctate, the punctures confused.
Antennae approximated basally; pronotum subquadrate, [79] Anthribidae: Anthribinae
distinctly narrower than elytra; lateral margin of pronotum Fig. 245 Sharpius venustus (Broun)
and medial margins of elytra subparallel; elytra with distinct A moderately small, elongate-oval fungus weevil 3.1-3.4
shoulders; legs robust; metafemora swollen. A South mm long, pubescent and scaled; integument black with
American species introduced into New Zealand from Aus- brilliant cream and orange markings. Head with rostrum
tralia for biological control of alligator weed (Alternanthera longer than wide, expanded apically; antennae straight,
phylloxeroides), and now well established in Northland. with a 3-segmented club, the basal segment symmetrical
and segment 8 elongate and slightly narrowed apically;
[77] Chrysomelidae: Galerucinae (Galerucini) pronotum slightly longer or slightly shorter than wide, with
Fig. 241 Adoxia vulgaris (Broun) transverse carina slightly curved and lateral carina short;
A medium-sized beetle 3.0-6.0 mm long, with head, pro- pygidium rugose in male and puncto-asperate in female.
notum, and elytra uniformly yellowish rust-brown, or head Beaten from various native and introduced shrubs. Widely
brown, pronotum orange-yellow, and elytra olive or yellow- distributed on both main islands, from Rotoehu State
ish brown with dark brown edges; appendages usually dark Forest to Invercargill.
brown. Upper body moderately convex, glabrous or nearly
so, densely punctate, the punctures fine; head and [79] Anthribidae: Choraginae
pronotum distinctly narrower than elytra. Collected from Fig. 246 Dysnocryptus pallidus Broun
rotten branches (e.g., Podocarpus), inflorescences of A small, oval, pubescent fungus weevil 1.2-2.3 mm long,
rangiora (Brachyglottis repanda), and beaten from shrubs. variegated reddish-brown and yellow, often with black or
Widely distributed across the North I. and northern South I. dark brown markings, rarely entirely black. Head with a
short, broad rostrum; antennae straight, with a 3-seg-

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rented club, the basal segment asymmetrical; pronotum brown or brown to almost black with sparse, moderately
wider than Iong, with transverse basal carina terminating long, yellowish-grey pubescence. Head and pronotum with
abruptly at lateral margin, and lateral carina absent; elytra meshed microsculpture; rostrum elongate, narrow; anten-
elongate, with striae containing large punctures. Adults nae slightly geniculate, inserted near middle of rostrum,
have been collected from leaf litter and beaten from dead with scape almost as long as remainder of antenna and
honeysuckle. Distributed in the southern part of the North I. club 3-segmented; pronotum narrower than elytra, slightly
and northeastern South I., including some offshore islands elongate and contracted anteriorly; elytra oval in outline,
(Stephens I., D'Urville l., and Chetwode Is). with coarsely punctate striae and convex interstices. Adults
occur on foliage of pohutukawa (Metrosideros excelsa),
[80] Belidae: Belinae and larvae live subcortically in branchlets. Recorded from
Fig. 247 Cyrotyphus tridens (Fabricius) Northland to Auckland and the Coromandel area, including
Alarge,ir gularyshapedwevil10-3m long,pale The Aldermen, Mercury Is, Little Barrier I., Ohena I., Hen
brown, mostly covered with whitish and pink scales. Head and Chickens Is, Coppermine I., and the Mokohinau Is.
and pronotum narrower than elytra and tapering anteriorly;
head not constricted behind eyes, with rostrum moderately [82] Curculionidae: Brachycerinae
elongate, broad; eyes prominent; antennae straight, with- Fig. 251 Anagotus turbotti (Spiller)
out a club; pronotum trapezoidal; elytra broad, tuberculate, A large weevil up to 24 mm long, brown or reddish brown
narrowed subapically; tarsal segment 3 strongly lobed. with whitish dorsal spots, well marked on middle and
Adults are frequently found in dead wood of many kinds. posterior elytra; ventral surface whitish. Elytta with large,
Recorded from the North I. and northern South I. cone-shaped posterior protuberances. Adults may be
found in the branches of native trees (e.g., ngaio), where
[81] Belidae: Aglycyderinae larvae are most likely wood-borers. Surviving only on rat-
Fig. 248 Aralius wollastoni (Sharp) free offshore islands such as the Three Kings, Poor
A moderately small, elongate, subparallel weevil 3.0-3.2 Knights, and Muriwhenau in the Hen and Chickens group.
mm long, dull brown, pubescent. Head abruptly constricted
behind eyes, with abrupt posterior angles; rostrum short, [82] Curculionidae: Brachycerinae
broad, shorter in females than in males; antennae straight, Fig. 252 Mandalotus miricollis (Broun)
with a distinct 2-segmented club; pronotum approximately A medium-sized, narrowly oval weevil 4.0-7.0 mm long,
subquadrate; elytra striate-punctate; tarsi with 2nd seg- matt brownish-grey, usually variegated with inconspicuous,
ment strongly lobed. Adults and larvae may be found small, irregularly shaped paler spots. Pronotum and
beneath bark of dead or dying branches of five-finger. particularly elytra with scattered elongate scales; rostrum
Apparently widely distributed. broad, short; antennae strongly elbowed; pronotum as
broad as elytra, with more or less distinct transverse
[81] Brentidae: Brentinae wrinkles; elytra with longitudinal shallow ridges. Occurs on
Fig. 249 Lasiorhynchus barbicornis ( Fabricius) sandy beaches and coastal vegetation, though occasion-
A very large, extremely nar owly elongate weevil 18-75 ally found in gardens in clusters of flax. Recorded in the
mm Iong (male; female 18-47 mm), punctate, with larger Auckland area.
punctures on elytra; minute yellowish pubescence on head,
pronotum, and elytra and additional brown and longer [82] Curculionidae: Curculioninae
pubescence on rostrum, antennae, and tarsi. Body matt Fig. 253 Andracalles spurcus (Broun)
brown variegated with reddish-brown spots, in some A small, oval weevil 1.9-2.4 mm long, matt brown, with
specimens almost uniformly dark brown; rostrum extremely brown and paler brown projecting scales. Rostrum narrow,
elongate, shorter in female than in male; antennae straight, long; antennae distinctly geniculate; pronotum slightly
insertdapclym,edianfls;protum narrower than elytra; elytra with scales forming irregular
with a median sulcus; elytra constricted subapically in rows; legs stout, with projecting scales. Adults have been
males, evenly tapering in females; tarsi long. Oviposition collected by beating Astelia sp. at dusk; also found on
sites are prepared by females from October to March by pohutukawa and in bryophytes in dense forest. Recorded
chewing a cavity in the bark of dying or suppressed trees from Northland, Auckland, and the Bay of Plenty.
(kauri, pigeonwood, rewarewa, tawa, karaka) or logs.
Newly hatched larvae bore radially into the wood, and [82] Curculionidae: Curculionidae
enlarge the tunnel as they grow. Ambrosia fungi and yeast Fig. 254 Myrtonymus zelandicus Kuschel
growing in the tunnels most likely serve as larval food. A minute, narrowly elongate weevil 0.7-0.8 mm long, pale
Recorded from the North I. and northern South I., mostly at yellowish or reddish brown with appendages usually a
lower altitudes. shade paler. Rostrum moderately elongate, sparsely punc-
tate; eyes completely reduced; pronotum longer than wide,
[81] Brentidae: Apioninae variably and sparsely punctate; elytra flattened, with fine
Fig. 250 Neocyba metrosideros (Broun) punctation forming obsolescent striae. As far as is known,
A small, pyriform weevil 1.8-2.5 mm long, glossy reddish this blind species is the smallest weevil in the world.

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Consistently found around the fine rootlets of all native [82] Curculionidae: Scolytinae
species of Myrtaceae, it occurs in the top 10 cm of soil Fig. 258 Hylastes ater (Paykull)
during wetter conditions and at greater depth, 30-60 cm, A medium-sized, narrowly elongate weevil 3.5-6.0 mm
during dry periods. Recorded from the northern North I. long, dark brown to black, rarely reddish brown; antennae
and tarsi usually reddish brown. Pronotum elongate and
[82] Curculionidae: Curculioninae densely punctate, the punctures round; elytral punctures
Fig. 255 Stephanorhynchus lawsoni Sharp round, and forming striae, with short inconspicuous
A small, distinctive weevil 3.2-4.8 mm long, characterised pubescence. Found mainly in industrial timber and under
by its long, deeply constricted head, conical prothorax, bark of Pinus radiata logs. Recorded from both main
tuberculate elytra, and extraordinarily long hind legs. These islands, ranging from Waipoua State Forest to Otago.
have strongly curved tibiae and femora with a large,
rectangular tooth, enabling the weevil to clasp bunches of [82] Curculionidae: Scolytinae
flower stamens and rake pollen towards the mouth. Fig. 259 Phloeosinus cupressi Hopkins
Common in Northland and the northern South I. A small, moderately short weevil 2.8-3.0 mm long, sub-
cylindrical with forebody slightly tapering anteriorly,
[82] Curculionidae: Cossoninae subglabrous with inconspicuous short and erect setae,
Fig. 256 Macroscytalus remotus (Sharp) moderately glossy. Head largely concealed by pronotum
A small, narrowly elongate weevil 2.5-3.0 mm long, glossy from above; rostrum short and broad; pronotum narrowing
yellowish red or dark reddish brown; subglabrous, with anteriorly, as broad as elytra at base; elytra mostly sub-
inconspicuous short and erect setae. Rostrum long, broad, patallel, with deep, coarsely punctate striae. An adventive
slightly expanding apically; antennae strongly geniculate; species recorded from Auckland on Cupressus torulosa.
pronotum almost as broad as elytra, rounded laterally;
elytra mostly subparallel, with angular basal angles and [82] Curculionidae: Platypodinae
with distinct punctate striae. Beaten from various trees, and Fig. 260 Platypus apicalis White
reared from larvae in Coprosma macrocarpa. First reported A medium-sized, subcylindrical, narrowly elongate weevil
from Auckland. 6-7 mm long, subparallel-sided, glossy reddish brown to
dark brown. Rostrum very short; antennae weakly geni-
[82] Curculionidae: Cossoninae culate, with a broad club; pronotum slightly elongate,
Fig. 257 Xenocnema spinipes Wollaston rectangular, shallowly emarginated mediolaterally; elytra
A somewhat flattened and elongate weevil 3.0-4.8 mm subparallel, finely striate; tarsi long and slender, with basal
long, with a short, broad rostrum, elongate prothorax, elytra segment strongly elongate. Adults are fully winged, and
with well marked striae and punctation, and heavy legs with may be collected using Malaise traps. They bore into a wide
strong teeth at the apex of the tibiae. Adults and larvae con- range of native and exotic trees, making tunnels about 2
struct galleries under loose bark of Agathis and Araucaria mm in diameter at right angles to the grain. These become
species. Restricted in distribution by the availability of host infected with ambrosia fungi, on which the larvae feed.
trees, and hence recorded mostly from northern localities. Occurs widely on both main islands and on the Chatham Is.

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 APPENDIX 1: Endangered beetle species APPENDIX 2: Major entomological collections
[modified after Molloy et al. 1994] in New Zealand
A number of species are as yet undescribed, and are AMNZ Auckland Institute & Museum
referred to by informal names or by `sp.' Synonyms and Private Bag 92-018, Auckland
superseded names are in parentheses. Attn: Mr John W. Early, Curator of Entomology
Tel: +64 9 309 0443 Fax: +64 9 379 9956
• Seven species are listed in Category A (highest priority
Internet: early@akmus.org.nz
threatened species):
Geodorcus (=Dorcus) auriculatus, G. ithaginis, G. 'Moe- CMNZ Canterbury Museum
hau', Mecodema costellum costellum, M. laeviceps, Pro- Rolleston Ave, Christchurch
dontria bicolorata, P. lewisi Attn: Simon Pollard, Curator of Invertebrate
• Ten species are listed in Category B (second priority Zoology
Tel:threatened+64
species): 3 366 8379 Fax: +64 3 366 8379
Internet: spollard@cantmus.govt.nz
Amychus granulatus, Anagotus stephenensis, A. turbotti,
Hadramphus spinipennis, Heterexis seticostatus, Lypero-
FRNZ Forest Research Institute
bius huttoni, Megadromus 1 sp., Oregus inaequalis, Pro-
Private Bag 3020, Rotorua
dontria grandis, Zecillenus (=Cillenum) tillyardi
Attn: Roger Crabtree, Curator of Entomology
Tel:
• Six species+64 7C (third
are listed in category 347 priority 5899 Fax: +64 7 347 5333
threatened species): Internet: crabtrer@fri.cri.nz
Amychus candezei, Anagotus fairburni, Hadramphus
stilbocarpae, Mecodema chiltoni, Oclandius laeviusculus, LUNZ Entomology Research Museum
Prodontria modesta Dept of Entomology & Animal Ecology, P.O. Box 84,
Lincoln University, Canterbury
• Ten species are listed in Category Χ (species which have Attn: Mr John W. M. Marris, Curator
Tel: +64
not been sighted 3
for a number of 325
years but which may 2811 Fax: +64 3 325 3844
still exist): Internet: marris @tui.lincoln.ac.nz
Hadramphus tuberculatus, Mecodema costellum `spelles',
M. punctellum, Megacolabus sculpturatus, Megadromus MONZ Museum of New Zealand Te Papa Tongarewa
antarcticus 2 subspp., Pericoptus nitidulus, Prodontria Natural Environment Dept, P.O. Box 467, Wellington
`Five Rivers', Stethaspis convexa, Thotmus halli Attn: Mr Ricardo L. Palma, Curator, Terrestrial
Arthropods
• Almost eighty
Tel: +64 species are assigned
4 to Category Ι
381 7361 Fax: +64 4 381 7310
(species about which little information exists, but Internet: ricardop@tepapa.govt.nz
which are considered threatened):
Amychus 1 sp., Blosyropus spinosus, Brullea antarctica, NZAC New Zealand Arthropod Collection
carabid `Kamo', Euconnus microcilipes, E. paracilipes, Landcare Research, Mount Albert Research Centre,
Holcaspis falcis, Lyperobius carinatus, Maorinus hunu- Private Bag 92-170, Auckland
aeformis + 1 sp., Maoripamborus fairburni + 1 sp., Meco- Attn: Dr Trevor K. Crosby, Curator
dema allani,+64
Tel: M. angustulum, M. 9atrox, M.849
brittoni, Μ. 3660 Fax: +64 9 849 7093
costellum lewisi, M. costellum obesum, M. curvidens, Μ. Internet: crosbyt@landcare.cri.nz
dunense, M. dux, M. howitti, M. integratum, M. litoreum,
M. minax, M. morio, M. nitidum, M. pavidum, Μ. pluto, Μ. OMNZ Otago Museum
proximum, M. pulchellum, Μ. quoinense, Μ. rex, Μ. P.O. Box 6202, Dunedin
strictum, Megacolabus bifurcatus, M. obesus, Megadro- Attn: Mr Anthony C. Harris, Curator of Zoology
mus capito (=bucolicus),
Tel: +64 M. compressus,
3 M. 477fultoni, Μ. 2372 Fax: +64 3 477 5993
haplopus, M. `Omarama', M. virens + 5 spp., Metablax 1
sρ., Mimopeus parallelus, Nesoptychias simpliceps, Note. New Zealand beetles may be found in institutional
Neuraphaconnus toronouii, Nothaldonus peacei, and private collections around the world. For instance, the
Oclandius cinnaraeus, Paralissotes mangonuiensis, Broun Collection with its many primary type specimens is
Paratorchus alifer, P. flexuosus, Pericoptus frontalis, held at the Natural History Museum, London, and the
Pheloneis gratiosus, Prodontria patricki, P. regalis, P. Field Museum, Chicago, has extensive holdings of Staph-
setosa, P. `Twizel', Sciacharis yakasensis, Stephano- ylinidae and Leiodidae from New Zealand. Despite name
rhynchus insolitus, Syrphetodes 1 sp., Tangarona pen- changes affecting some institutions, the list of repositories
sus, Tychanopais tuberosus, Zeopoecilus 5 or 6 spp. published by Watt (1979c) is still a useful introduction.

—184—
 APPENDIX 3
Key to tribes of Carabidae in New Zealand
[provided by J.F. Lawrence]

1 Clypeus broader than distance between antennal sock- - Elytra with an inner longitudinal ridge, its end visible
ets; eyes large and protruding; protibia with 2 spurs beneath edge of elytron, level with apical end of
terminal; lateral pronotal carinae absent or incomplete epipleuron; head with frontal furrows not extending
posteriorly ... Cicindelini behind eyes ... 10
—Clypeus narrower than distance between antennal
sockets; without other characters in combination ... 2 10(9) Maxillary palp with penultimate segment setose;
male protarsi with 2 segments dilated and dentate on
2(1) Metepimeron not visible between posterior edge of inner face ... Zolini
metepisternum and anterior edge of 1st ventrite ... 3 —Maxillary palp with penultimate segment glabrous;
—Metepimeron visible as a lobe between metepisternum male protarsi unmodified, or only slightly so
and 1st ventrite ... 5 Psydrini

3(2) Procoxal cavities closed behind; apical segment of 11(7) Elytra with apex rounded as a single curve, or
maxillary palp slender and fusiform; length less than acutely angled at apex; Abdominal apex usually
15 mm ... Migadopini concealed from above; cloration rarely yellow and
—Procoxal cavities open behind; apical segment of black ... 12
maxillary palp more or less expanded and truncate —Elytra with apex transversely or obliquely truncate,
apically; length more than 15 mm ... 4 exposing abdominal apex from above; coloration
often yellow and black; body often flattened, or with
4(3) Mandibles multidentate, smooth on upper surfaces; prothorax elongate and narrow ... 15
clypeus without setiferous punctures ...Cychrini
—Mandibles not toothed, rugose on upper surfaces; 12(11) Head with a single pair of supraorbital setiferous
clypeus with a setiferous puncture at each anterior punctures ... Harpalini
angle ... Carabini —Head with 2 pairs of supraorbital setiferous punctures
... 13
5(2) Body pedunculate, with a distinct, narrow waist or
peduncle between prothorax and elytra; scutellum 13(12) Clypeus emarginate, with a pale membrane bet-
entirely contained within peduncle, not forming a ween clypeus and labrum (usually within emargination);
wedge between elytral bases ... 6 mandibles characteristically bifurcate, with a large
—Body not pedunculate, or if slightly so then scutellum mesal tooth; labrum often deeply emarginate
extending behind peduncle and forming a wedge Licinini
between elytral bases ... 7 —Clypeus not emarginate, without an obvious pale
membrane between it and labrum ... 14
6(5) Mesocoxal cavities open laterally, partly closed by
mesepimeron ... Clivinini 14(13) Elytra with an inner longitudinal carina, its apical
—Mesocoxal cavities closed laterally by meeting of end visible below lateral edge towards apex, causing
sterna ... Broscini end of epipleuron to appear twisted; carina sometimes
concealed by lateral edge of abdomen, which fits into
7(5) Mandibles with a setiferous puncture in concavity a groove between it and epipleural apex; mentum
(scrobe) along outer edge ... 8 usually bifid at apex ... Pterostichini
—Mandibles without a setiferous puncture in concavity — Elytra without an inner carina or, if present, not ex-
(scrobe) along outer edge ... 11 tending to lateral edge; mentum usually simple or with
a single median apical tooth ... Agonini
8(7) Maxillary palp with terminal segment very small
Bembidiini 15(11) Mentum supported on a projecting submentum;
—Maxillary palp with terminal segment usually about as head not sharply constricted behind eyes; mandibles
long as penultimate segment ... 9 with hollow outer face; claws often pectinate
...Lebiini
9(8) Elytra without an inner longitudinal ridge beneath —Without a projecting submentum; head sharply
apical edge; head with frontal furrows extending constricted behind eyes; mandibles without hollow
beyond posterior edge of eyes ... Trechini outer face; claws simple ... Pentagonicini

—185-
 APPENDIX 4: Distribution of collection records for beetle families in New Zealand
(Nth, North I.; Sth, South I.; Stw, Stewart I.; Ke, Kermadec Is; 3K, Three Kings Is; Ch, Chatham
Is; Sn, The Snares Is; An, Antipodes Is; Bo, Bounty Island; Au, Auckland Is; Ca, Campbell I.).

Family Nth Sth Stw Ke 3Κ Ch Sn An Bo Au Ca

Aderidae •
Agyrtidae •
Anobiidae • • • • • •
Anthicidae •
Anthribidae • •
Archeocrypticidae • •
Belidae •
Bostrichidae •
Bothrideridae •
Brentidae •
Buprestidae •
Byrrhidae •
Cantharidae •
Carabidae •
Cavognathidae • •
Cerambycidae •
Cerylonidae • •
Chaetosomatidae •
Chalcodryidae •
Chelonariidae
Chrysomelidae • •
Ciidae •
Clambidae •
Cleridae •
Coccinellidae •
Colydiidae •
Corticariidae •
Corylophidae •
Cryptophagidae •
Cucujidae •
Curculionidae • •
Dermestidae •
Derodontidae •
Dryopidae •
Dytiscidae •
Elateridae •
Elmidae •
Endomychidae •
Erotylidae •

—186—
Family Nth Sth Stw Ke 3Κ Ch Sn An Bo Au Ca
Eucinetidae
Eucnemidae • .

Gyrinidae .9

Heteroceridae
Histeridae
Hydraenidae
Hydrophilidae
Jacobsoniidae
Laemophloeidae . •

Languriidae • •

Leiodidae
Limnichidae
Lucanidae • •
Lycidae
Melandryidae
Melyridae
Monotomidae
Mordellidae •

Mycetophagidae
Nemonychidae
Nitidulidae
Nosodendridae
Oedemeridae
Phalacridae
Phloeostichidae
Phycosecidae • . 9

Prostomidae
Ptiliidae
Ptilodactylidae
Pyrochroidae
Rhipiphoridae
Rhysodidae .9

Salpingidae
Scarabaeidae
Scirtidae
Scraptiidae
Scydmaenidae
Silvanidae
Staphylinidae
Tenebrionidae
Trogidae
Trogossitidae
Ulodidae
—187—
 TAXONOMIC INDEX
Only coleopteran taxa are indexed, since other organisms mentioned in the text, e.g., host taxa, are
largely derived from other published sources. Names are listed in a typographical hierarchy, as
follows. CAPITALS: taxa above family level. Bold: the 82 families that occur in New Zealand.
Normal: exotic families, family synonyms, subfamilies, tribes. I talic: genera, species. Page numbers
on which a taxon is keyed out are suffixed k'. Bold type denotes the page on which a family is
described, and Italic the page on which a taxon is il ustrated.

Abraeinae 24 Anthicus 63: hesperi 63, 159, 179

Acanthoscelides 106: obtectus 125 Anthreninae 40
Acrossidius tasmaniae 31, 139, 171 Anthrenocerus 40: australis 41
Acrotrichinae 25 Anthrenus 40: verbasci 40
Actizeta 60: albata 61, 157, 179 Anthribidae 16, 68, 99k, 106k, 125, 164, 186
Adeliini 61 Anthribinae 68, 69k, 164
ADEPHAGA 13, 15, 18, 20, 97k, 105k, 113, 114, 121, Aphodiinae 30, 31k, 139
122, 126 Aphtora 60
Aderidae 64, 101k, 109k, 160, 186 Apion 124
Adoxia 125: vulgaris 67, 163, 181 Apioninae 16, 69, 70k, 97, 106k, 165
Adrastia clarkei 26, 133 Aralius wollastoni 69, 125, 164, 182
Aegialitinae 63 Archaeoglenes 98: costipennis 117
Aesalinae 29, 137 Archeocrypticidae 13, 56, 102k, 109k, 155, 186
Agapythinae 13, 47k, 112k, 150 Archeocrypticus topali 56, 155, 177
Agapytho 101: foveicollis 47, 15, 175 Archeoglenes 60
Agasicles hygrophila 67, 162, 181 ARCHOSTEMATA 13, 15, 122
Agathinus tridens 124, 125 Arhopalus tristis 65
Aglycyderinae 69k, 97, 98, 106k, 164 Aridius nodifer 55
Agnosthaetus vicinus 27, 136, 170 Arnomus 125: brouni 67, 163, 181
Agonini 185 Arpediomimus kronei 127
Agrypninae 37, 144 Arthopus 59
Agyrtidae 12, 25, 103k, 111k, 132, 186 Arthracanthus 45
Ahasverus 48: advena 48 Arthrolips 54
Alema spatiosa 125 Artystona 61: erichsoni 117
Aleochara 16, 27: hammondi 27, 135, 169 Ascetoderes 52: obsoletus 52, 153, 177; paynteri 52
Aleocharinae 16, 27, 28k, 135 Asilis 38: fulvithorax 38, 145, 174; fumida 124
Alleculinae 60, 61, 126 Asphaerites 26
Allocharis 125 Atomariinae 50
Allocinops brookesi 58, 126 Attageninae 40
Alphitobius 61 Attagenus 40
Alticini 66, 67, 162
Amarygmus 61 Baeocera scutellaris 27, 126, 135, 170
Amplectopus pallicornis 32, 141, 172 Belidae 69, 97, 98, 99k, 106k, 124, 125, 164, 186
Amychus 184: candezei 37, 144, 173, Belinae 69k, 106k, 124, 164
184; granulatus 184 Bembidiini 185
Anagotus 71: fairburni 125, 184; Bembidion 21
stephenensis 184; turbotti 70, 165, 182, 184 Blosyropus spinosus 184
Anchytarsinae 35, 143 Bostrichidae 13, 41, 42, 44, 98k, 103k, 111k, 118, 124,
Andracalles spurcus 70, 166, 182 146, 186
Anisomeristes 54: sharpi 55, 154, 177 BOSTRICHIFORMIA 38
Anobiidae 16, 17, 41, 42, 44, 102k, 107, 108k, 111k, Bostrichinae 41
117, 124, 147, 186 BOSTRICHOIDEA 39, 98k, 100k, 102k, 104k
Anobiinae 42, 108k, 147 Bothrideridae 15, 52, 105k, 110k, 111k, 153, 186
Anobium: punctatum 42; ruficorne 42 Bothriderinae 52, 153
Anommatinae 52, 111k, 153 Brachycerinae 70, 71k, 165
Anommatus 52: duodecimstriatus 52, 153, 177 Brachypeplus 46
Antarcticodomus 63 Brentidae 69, 97, 99k, 106k, 124, 125, 165, 186
Anthicidae 63, 64, 101k, 109k, 159, 160, 186 Brentinae 70k, 106k, 165
Anthicinae 63, 159 Brontinae 48

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Brontopriscus pleuralis 48, 150, 176 Chrysomelidae 12, 13, 16, 17, 54, 60, 66, 97, 99k,
Broscini 185 100k, 106, 125, 127, 162, 163, 186
Brounia 35, 98k: thoracica 14, 36, 143, 173 Chrysomelinae 66, 67k, 162
Brouniphylax squamiger 59 CHRYSOMELOIDEA 65, 98k, 99k, 106k, 125
Bruchidius villosus 67, 162, 181 Chrysopeplus 61
Bruchinae 16, 66, 67k, 98, 100k, 106k, 162 Cicindelidae 21
Brullea antarctica 184 Cicindelinae 21k, 106k, 128
Buprestidae 12, 16, 33, 44, 97k, 107k, 120, 124, 142, Cicindelini 185
186 Ciidae 16, 56, 105k, 110k, 117, 155, 186
Buprestinae 33 Ciinae 57, 155
Buprestis aurulenta 33 Cillaeinae 46
BUPRESTOIDEA 33 Cis zeelandicus 57, 117, 155, 178
Byrrhidae 12, 13, 33, 40, 90k, 107, 108k, 123, 142, 186 Clambidae 32, 104k, 108k, 141, 186
Byrrhinae 33, 142 Clambus 32: domesticus 32, 141, 172
BYRRHOIDEA 33, 99k, 100k, 104k, 106k, 107k CLAVICORNIA 46
Byrrocryptus 35: urquharti 36, 143, 173 Cleridae 13, 16, 44, 101k, 101k, 112k, 124, 148, 186
Clerinae 44
Cacephatus incertus 125 CLEROIDEA 43, 102k
Caccomolpus 125 Clivina basalis 21, 129, 168
Cafius litoreus 28, 137, 171 Clivinini 185
Camiarinae 25, 26k, 132 Coccidulinae 54
Cantharidae 16, 38, 100k, 101k, 107k, 118, 124, 145, Coccinella 117: leonina 54, 125, 154, 177
186 Coccinellidae 15, 16, 54, 55, 104k, 110k, 117, 125,
Carabidae 13, 16, 21, 60, 97k, 106k, 113, 114, 119, 126, 154, 186
126, 128, 129, 185, 186 Coccinellinae 54, 154
Carabinae 21, 22k, 128 Coelometopinae 60
Carabini 185 Colon 26: hirtale 26, 133
CARABOIDEA 20 Coloninae 25, 26k, 133
Carabus 21 Colydiidae 12, 13, 16, 52, 53, 58, 104, 105k, 110k,
Cardiophorinae 37 111k, 120, 124, 156, 186
Carpelimus 27, 136, 170 Colydiinae 59, 156
Carpophilinae 46 Colymbetinae 22
Carpophilus 46 Copelatinae 22
Cathartocryptus maculosus 51 Coprostygnus 28
Catopsolius 26 Corticaria 55
Cavognathidae 12, 16, 50, 103k, 112k, 127, 152, 186 Corticariidae 13, 16, 46, 55, 105k, 111k, 154, 186
Cerambycidae 13, 16, 17, 37, 44, 58, 65, 99k, 106k, Corticariinae 55
125, 161, 162, 186 Corylophidae 54, 104k, 110k, 154, 186
Cerambycinae 65k, 161 Corylophinae 55
Ceratognathus: gibbosus 29; parrianus 30, 98, 137, Cossoninae 70, 71k, 166, 167
171; passaliformis 29 Costelytra 98: zealandica 30, 31, 127, 140, 172
Cerodolus 61 Cotes 63: crispi 63, 160, 180
Cerylidae 52 Creophilus oculatus 116
Cerylonidae 52, 105k, 110k, 153, 186 Cryptarchinae 46
Ceryloninae 53 Cryptamorpha brevicornis 48
Chaerodes 60: trachyscelides 61, 157, 178 Cryptocephalinae 66, 67k, 99, 163
Chaetosomatidae 12, 44, 102k, 112k, 148, 186 Cryptodacne 52: brouni 120
Chaetosoma 44: scaritides 44, 148, 175 Cryptolestes 49: ferrugineus 49; pusilloides 49
Chaetosomodes 44 Cryptophagidae 16, 50, 51, 103k, 105, 110k, 112k, 115,
Chalcodrya 60: variegata 60, 157, 178 152, 186
Chalcodryidae 13, 16, 59, 102k, 109k, 157, 186 Cryptophaginae 50, 51, 152
Chelagyrtodes 26 Cryptophagus 50
Chelonariidae 14, 35, 36, 98k, 106k, 118, 143, 186 Cryptorhynchini 71
Chelonarium 36 Cryptosomatulini 51
Chilocorinae 54 Ctenicera 37
Cholevinae 25, 26k Ctenognathus novaezelandiae 21, 129, 168
Choraginae 68, 69k, 164 Cucujidae 14, 16, 48, 49, 101k, 103k, 105, 109k, 112k,
Chrysolina: hyperici 67, 162, 181; quadrigemina 67 120, 126, 151, 186

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 CUCUJIFORMIA 38, 42 Epichorius longulus 123
CUCUJOIDEA 46, 99k, 101k, 102k, 104k, 125 Epurea 46
Curculionidae 13, 16, 17, 27, 57, 70, 99k, 106k, 116, Erotylidae 16, 51, 103k, 112k, 120, 152, 186
118, 125, 127, 165-167, 186 Euaesthetinae 27, 28k, 108, 136
Curculioninae 70, 71k, 166 Eualema speculifera 125
CURCULIONOIDEA 14, 67, 98k, 106k, 125 Eucinetidae 32, 102k, 108k, 141, 187
Curimus 107 Eucinetus stewarti 32, 141, 172
Cychrini 185 EUCINETOIDEA 100k, 102k, 104k, 108k
Cyclaxyra 50, 105, 110, 112: impressa 49, 151, 176; Eucnemidae 36, 100k, 107k, 144, 187
politula 49 Eucolaspis brunnea 67, 127, 163, 181
Cyclaxyrinae 13, 49, 151 Euconnus microcilipes 184
Cyphon 32 Euconnus paracilipes 184
Cyphonidae 31 Euderia 41, 98: squamosa 41, 124, 146, 174
Cyrotyphus 69: tridens 69, 164, 182 Euderinae 13, 41, 103, 146
Euglenidae 64, 101
Dacninae 52, 152 Eumolpinae 66, 67k, 163
Dascyllidae 31 Eupines 28
Dasytes 45: subcyaneus 45, 149, 175 Euplectopsis 28
Dasytidae 45, 101k, 112k Euxestinae 53, 153
Dasytinae 45, 149 Exapion ulicis 70
Demtrius 61 Exocalopus pectinatus 62, 159
Dendroblax earlii 30, 138, 171 Exochomus 54
Dendrophilinae 24
Denticollinae 37, 144 Galerucinae 66, 67k, 162, 163
Dermestes 40: maculatus 41, 123 Galerucini 66, 67, 163
Dermestidae 16, 17, 40, 100k, 103k, 107, 108k, 123, Geodorcus: auriculatus 30, 138, 171, 184; ithaginis 14,
146, 186 184; 'Moehau' 184
Dermestinae 40, 108k Gnathocerus 61
Derodontidae 14, 39, 100k, 108k, 145, 186 Gonocephalum 61
DERODONTOIDEA 38, 100k Grynoma regularis 43, 147, 175
Derolathrinae 111k Gymnusini 14
Derolathrus 39, 105 Gyrinidae 13, 14, 22, 97k, 106k, 130, 187
Diagrypnodes 63: wakefieldi 63, 159, 179 Gyrininae 22, 130
Diaperinae 60 Gyrinus: convexiusculus 22, 23, 130, 168; hutton 22
Dinoderinae 41, 146 Gyrophaena 27
Dinoderus 41: minutus 41, 146, 174
Discyllidae 31 Habrocerinae 27, 29k, 118, 135
Dorcatominae 42 Habrocerus capillaricornis 27, 135, 170
109 Doxzilra Hadramphus: spinipennis, stilbocarpae, tuberculatus
Dryocora 61: howitti 61, 158, 179 184
Dryophthorinae 70, 71k Hadrobregmus magnus 42, 147, 174
Dryopidae 34, 63, 99k, 107, 118, 142, 186 Halmus 54: chalybaeus 54
Dynastinae 30, 31k, 140 Hapalips 51: prolixus 51, 152, 176
Dysmorphocerinae 38, 145 Harpalinae 21, 22k, 129
Dysnocryptus pallidus 68, 164, 181 Harpalini 185
Dytiscidae 22, 97k, 106k, 126, 129, 130, 186 Helodidae 31
Dytiscinae 22 Heterexis seticostatus 184
Heteroceridae 35, 105k, 107k, 143, 187
Elateridae 13, 17, 37, 100k, 107k, 123, 144 Heterocerinae 35, 143
ELATERIFORMIA 31 Heterocerus novaeselandiae 35, 143, 173
Elaterinae 37 HETEROMERA 56
ELATEROIDEA 36, 99k, 100k, 107k Heteronychus arator 31, 140, 172
Elmidae 34, 63, 99k, 108k, 142, 186 Hippodamia 54
Enarsus bakewelli 120 Histerldae 23, 98k, 107k, 131, 187
Endomychidae 53, 55, 105k, 110k, 111k, 153, 154, 186 Histerinae 24
Enicmus caviceps 55, 154, 177 Hoherius 68
Enochrus tritus 23, 130, 168 Holcaspis falcis 184
Enopliinae 44, 148 Holoparamecinae 53, 105k, 111k, 154

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Holoparamecus 53, 55: tenuis 54, 154, 177 Lithostygnus 55, 105
Horelophinae 13, 23, 130 Loberonotha 51
Horelophus 14, 23: walkeri 23, 130, 168 Loberus: depressus 51, 127; nitens 51
Hydora picea 34, 142, 173 Longitarsus jacobaeae 67
Hydraena 24 Lophocaterinae 43, 44k, 112k, 147
Hydraenidae 12, 13, 24, 99k, 108k, 131, 132, 187 Lorelus 61
Hydraeninae 24, 131 Loxomerus nebrioides 21, 126, 128, 168
Hydrophilidae 13, 14, 23, 24, 99k, 108k, 118, 120, 130, Lucanidae 16, 29, 98k, 106k, 118, 137, 138, 187
131, 187 Lucaninae 29, 137, 138, 171
Hydrophilinae 23, 130 Lycidae 13, 16, 37, 100k, 107k, 118, 124, 144, 187
HYDROPHILOIDEA 23, 99k Lyctinae 41, 103, 111, 146
Hydroporinae 22, 129, 130 Lyctus: brunneus 41, 146, 174; linearis 41; planicollis
Hylastes ater 71, 167, 183 41
Hylobia: nubeculosa 57, 155, 178; pulla 126 Lyperobius: carinatus 184; huttoni 184
Hyperomma 28
Hyphalinae 34, 107k Macratria 63: exilis 63, 160, 180
Hyphalus 34, 99 Macratriinae 63, 160
Hypodacne 53 Macraulacinae 36
Hypodacnella 52: rubriceps 53, 153, 177 Macroscytalus remotus 70, 166, 183
Malthininae 38
Illeis 54 Mandalotus miricollis 70, 165, 182
Inocatops elongellus 26, 132 Maoraxia eremita 33
Inopeplinae 62, 63, 100k, 102, 108, 109k, 159 Maorinus hunuaeformis 184
Maoripamborus fairburn 21, 119, 128, 168, 184
Jacobsoniidae 12, 39, 103k, 105k, 111k, 145, 187 Mecodema 21: allani, angustulum, atrox, brittoni, chil-
toni, costellum costellum, costellum lewisi, costellum
Korynetinae 44 obesum, costellum 'spelles', curvidens, dunense, dux,
Kuschelydrus 97: phreaticus 22, 130, 168 howitti, integratum, laeviceps, litoreum, minax, morio,
nitidum, pavidum, pluto, proximum, pulchellum, punc-
Laemophloeidae 47, 49, 101k, 103k, 105, 109k, 151, tellum, quoinense, rex, strictum 184
187 Medon zeelandicus 27, 137, 171
Lagriinae 60, 61, 157 Megacolabus: bifurcatus, obesus, sculpturatus 184
Lagrioida 63: brouni 63, 160, 180 Megadromus 21, 184: antarcticus 21, 184; bucolicus,
Lagrioidinae 63, 109k, 160 capito, compressus, fultoni, haplopus, `Omarama',
Lamiinae 65k, 161 virens 184
Lampriminae 29, 138 Megatominae 40, 146
Lancetes lancelatus 126 Melandryidae 12, 13, 57, 64, 102k, 109k, 126, 155, 187
Languriidae 51, 103k, 112k, 127, 152, 186 Melandryinae 57, 155
Larainae 34, 142 Melanophthalma 55
Laricobiinae 39, 145 Melasinae 36, 144
Lasiorhynchus barbicornis 70, 125, 182 Melolonthinae 30, 31k, 139, 140
Lathridiidae 13, 46, 55, 105, 111 Melyridae 45, 101k, 112k, 149, 187
Lathridiinae 55, 154 Menimus 60
Latridiidae 55 Meropathus 24, 97, 99, 108
Lebiini 185 Meropathus zelandicus 24, 132, 169
Leiodidae 12, 13, 25, 103k, 111k, 118, 132, 133, 187 Merophysiidae 53
Leiodinae 25, 26k Merophysiinae 53
Lemodinae 63, 160 Metablax 184
Lenax 46, 100, 103, 104, 111: mirandus 14, 46, 47, 149, Metacorneolabium 12
175 Metasclera 64
Lepidopteryx 43: nigrosparsa 43, 148, 175 Metaxina 44, 101, 111: ornata 45, 148, 175
Licinini 185 Metophthalmus 55
Limnichidae 35, 99k, 107k, 143, 187 Metriorrhynchus erraticus 38
Limnichinae 35, 108k, 143 Microbrontes 49
Limnichus 35: nigripes 35, 143, 173 Microbrontes lineatus 49, 151, 176
Liochoria huttoni 34, 142, 173 Microsilpha 14, 28: litorea 27, 133, 169
Liodessus plicatus 22, 129, 168 Microsilphinae 27, 28k, 100k, 108, 133
Lissominae 37 Migadopinae 21, 22k, 128

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Migadopini 185 Orthoperus 54, 110
Mimopeus 61: elongatus 61, 158, 179; opaculus 60; Oryzaephilus 48: surinamensis 48, 151, 176
parallelus 184; subcostatus 119 Osoriinae 27, 28k, 136
Monommatidae 59 Othius 28
Monotoma 46, 100, 103, 104, 110: spinicollis 47, 150, Oxytelinae 27, 28k, 136
175
Monotomidae 14, 46, 100k, 103k, 104k, 105, 110k, Pachyurinus 69
111k, 149, 150, 187 Paederinae 27, 29k, 137
Monotominae 47, 149, 150 Paralissotes mangonuiensis 184: reticulatus 30, 137,
Mordella antarctica 58, 156, 178 171
Mordellidae 57, 102k, 109k, 156, 187 Paramulinae 55
Mordellinae 58, 156 Parasiagonum hudsoni 27, 135, 170
Mycetaea 53: lucidus 53; subter-ranea 54, 153, 177 Paratorchus 28: alifer 184; flexu-osus 184
Mycetaeinae 53, 110k, 153 Parepierus 24
Mycetophagidae 50, 56, 105k, 110k, 120, 155, 187 Parisopalpus maclearyi 127
Mycetophaginae 56, 155 Parnida agrestis 34, 142, 173: longulus 34
Myllaena 27 Paropsis charybdis 66
Myrtonymus zelandicus 70, 166, 182 Partystona 61
MYXOPHAGA 13, 15 Paupris 44
Pelecatominae 58, 156
Nacerdinae 62 Peltidae 43
Nanosellinae 25 Pentagonicini 185
Nascioides enysi 33, 120, 124, 142, 172 Pericoptus 30: frontalis, nitidulus 184; truncatus 31,
Nausibius clavicornis 48 140, 172
Necrobia 44: ruficollis 44; rufipes 44 Perimylopidae 60
Necrophilus 25, 132: prolongatus 25, 132, 169 Phalacridae 13, 49, 97, 98k, 105, 110k, 112k, 151, 187
Nemonychidae 12, 16, 67, 99k, 106k, 125, 163, 187 Phalacrinae 49
Neocercus 50: electus 50, 127 Phalacrus 49: uniformis frigoricola 50
Neocharis simplex 37, 144, 173 Pheloneis gratiosus 184
Neocicindela tuberculata 21, 128, 168 Philothermus 52, 53
Neocyba 69, 97: metrosideros 70, 125, 165, 182 Phloeocharinae 27, 29k, 134, 169
Nesoptychias simpliceps 184 Phloeosinus cupressi 71, 167, 183
Neuraphaconnus toronouii 184 Phloeostichidae 13, 47, 101k, 102k, 104k, 105, 110k,
Nitidulidae 16, 46, 100k, 104k, 108, 111k, 118, 149, 112k, 123, 150, 187
187 Phreatodessus 22
Nitidulinae 46, 149 Phrenapatinae 110k
Nosodendridae 33, 40, 103k, 107, 108k, 145, 187 Phycochus graniceps 31, 139, 171
Nosodendron 33: ovatum 40; zealandicum 40, 145, 174 Phycosecidae 12, 45, 103k, 112k, 149, 187
Nothaldonus peacei 184 Phycosecis 45: limbata 45, 149, 175
Nothoderodontus 39: gourlayi 39, 145, 174 Phyllobaeninae 44
Nothotelus 64: usitatus 64, 126, 161, 180 Phymatophaea 44: lugubris 124; violacea 45, 148, 175
Notolaemus 49 PHYTOPHAGA 65
Notoptenidium lawsoni 25, 132, 169 Physobryaxis inflata 117
Nototorchus ferrugineus 27, 136, 170 Picrotus 50
Piestinae 27, 28k, 135
Ocalea 28 Pilipalpinae 12, 62, 158, 159
Ochthebiinae 24, 132 Pimeliinae 60, 61, 157
Oclandius cinnaraeus 184: laevi-usculus 184 Pityobiinae 37
Odontria giveni 31, 139, 171 Placonotus 49
Oedemeridae 61, 101k, 109k, 127, 158, 187 Platipidia 46: asperella 46, 149, 175
Oedemerinae 62, 158 Platisus 14, 48, 120, 126, 151, 176
Oemona hirta 58, 65, 161, 180 Platypodinae 70, 71k, 98, 106, 167
Oligota 28 Platypus 47: apicalis 71, 167, 183
Omaliinae 12, 14, 27, 28k, 100k, 133 Podaena latipalpis 131, 169
Omaliomimus albipennis 27, 133, 169 POLYPHAGA 12, 13, 15, 18, 22, 97k, 105k, 115, 121,
Omedes 61 122, 126
Onthophagus granulatus 31, 139, 171 Porrostoma rufipenne 38, 124, 144, 173
Oregus inaequalis 184 Priasilpha 102, 103: obscura 47, 123, 150, 176

-192-
Priasilphidae 47 Salpinginae 63, 159
Priasilphinae 47k, 112k, 150 Salpingus 63: bilunatus 63, 159, 179
Prioninae 65k, 161 Saphophagus 39: minutus 12, 39, 145, 174
Prionoplus 58: reticularis 65, 125, 161, 180 Saprininae 24, 131
Pristoderus antarcticus 59, 156, 178 Saprinus 24
Prodontria 30: bicolorata, 'Five Rivers', grandis 184; Saprosites 30
lewisi 30, 31, 140, 172, 184; modesta, setosa, Scaphidiinae 26-28k, 111k, 126, 135
patricki, regalis, 'Twizel' 184 Scarabaeidae 13, 16, 17, 30, 60, 98k, 106k, 127, 139,
Prostomidae 61, 104, 105k, 110k, 158, 187 140, 187
Proteininae 27, 29k, 100k, 134 SCARABAEIFORMlA 29
Protelaterinae 37 Scarabaeinae 30, 31k, 139
Protopeltinae 43, 44k, 112k SCARABAEOIDEA 15, 16, 29, 98k, 106k
Pselaphidae 13 Scaritinae 21, 22k, 129
Pselaphinae 12, 13, 16, 26-28k, 100k, 108k, 117, 134 Sciacharis yakasensis 184
Pseudhelops 61 Scirtidae 12, 13, 31,101k, 108k, 117, 141, 187
Pseudochelonarium 36 SCIRTOIDEA 31
Pseudophloeocharis australis 27, 134, 170 Scolytinae 41, 44, 57, 70, 71k, 98, 106, 167
Pseudopsinae 27, 29k, 136 Scopodes 21
Pseudopsis arrowi 27, 136, 170 Scraptildae 64, 101k, 109k, 126, 160, 187
Psydrini 185 Scraptiinae 64, 65
Pterostichini 185 Scraptogetus 64
Ptiliidae 12, 16, 24, 99, 104k, 107k, 132, 187 Scydmaenidae 12, 13, 26, 103k, 111k, 133, 187
Ptiliinae 25, 132 Scydmaeninae 26, 133
Ptilodactylidae 35, 100k, 107k, 118, 143, 187 Scymninae 54
Ptininae 16, 42, 102, 111k, 124, 147 Scymnus 54, 117
Ptinus: speciosus 42, 117, 124, 147, 174; tectus 42 Sepedophilus 27, 28, 134, 170
Pycnomerinae 59, 110k Sericoderinae 55, 154, 177
Pycnomerus sophorae 124 Sharpius venustus 68, 164, 181
Pyrochroidae 62, 101k, 109k, 127, 158, 159, 187 Silphidae 25
Pyronota festiva 30 Silphotelus 108: nitidus 27, 134, 170
Pythidae 62 Silvanidae 16, 47, 101k, 102k, 109k, 150, 151, 187
Silvaninae 48, 151
Quedius 28 Silvanus 48
Sirrhas 60
Reesa vespulae 40 Sitona discoidea 127
Reichardtia 24: pedatrix 24, 131, 169 Soronia 46
Rentonidium costiventris 43, 126, 147, 174 Sphaeridiinae 23, 131
Rentoniinae 43, 44k, 112k, 126, 147 Sphaerothorax 32
Rentonlini 43 Spondylinae 65k
Rhinorhynchinae 68, 163 Staphylinidae 12-14, 16, 26, 100k, 107k, 108k, 111k,
Rhinorhynchus rufulus 68, 125, 163, 181 116, 117, 118, 126, 127, 133-137, 187
Rhipiphoridae 58, 98k, 100k, 109k, 118, 126, 156, 187 STAPHYLlNIFORMIA 23
Rhipistena 58: lugubris 58, 156, 178 Staphylininae 27, 29k, 137
Rhizonium antiquum 59, 156, 178 STAPHYLINOIDEA 12, 15, 24, 99k, 100k, 102k, 104k
Rhizopertha 41: dominica 41 Stegobium paniceum 42
Rhizophaginae 104 Stephanorhynchus insolitus 184: lawsoni 70, 166, 183
RHYNCHOPHORA 67 Stethaspis 30: convexa 184
Rhynchophorinae 70, 71 Stylogymnusa subantarctica 12, 14
Rhysodidae 20, 97k, 106k, 118, 128, 187 Syncalyptinae 33
Rhysodini 20 Syndesinae 29
Rhyzobius 54 Symorthus insularis 123
Rhyzodiastes proprius 20, 128, 168 Syrphetodes 59, 184: marginatus 59, /57,178
Rodalia 54, 117
Rygmodus incertus 120: tibialis 23, 131, 169 Tachyporinae 27, 29k, 134
Rytinotus 58 Tangarona pensus 184
Tanychilus 61: sophorae 126
Sagola 28: laminata 27, 134, 170 Tarsosteninae 44
Salpingidae 62, 100k, 102k, 108, 109k, 110k, 159, 187 Techmessa: concolor 62, 158, 179; telephoroides 127

-193-
Techmessodes 62
Tenebrio 61
Tenebrionidae 13, 14, 16, 56, 59, 60, 98, 101k, 109k,
110k, 117, 119, 157, 158, 187
Tenebrioninae 60, 61, 158
TENEBRIONOIDEA 12, 15, 56, 98k, 100k, 101k, 104k
Tenebroides mauritanicus 43
Thallis polita 52, 152, 177
Thanoclerinae 44, 148
Thelyphassa lineata 62, 158, 179
Thoramus 37: wakefieldi 37, 144, 173
Thortus ovalis 51, 152, 176
Thotmus halli 184
Tomogenius latipes 24, 131, 169
Trechinae 21, 22k, 129
Trechini 185
Tribalinae 24
Tribolium 61
Trichelodes 40
Trichelodini 40
Trichocolposinus 63
Trinodinae 40
Triphyllus 56: hispidellus 56, 155, 177
Trogidae 13, 30, 98k, 106k, 118, 138, 187
Trogoderma 40: maestum 41, 146, 174
Trogossitidae 13, 16, 43, 104k, 112k, 126, 147, 148,
187
Trogossitinae 43k, 112k, 148
Trox scaber 30, 138, 171
Tychanopais tuberosus 184
Tymbopiptus valeas 12
Typhaea stercorea 120

Ulodidae 59, 60, 101, 102k, 109k, 119, 157, 187

Uloma 60
Ulomotypus 60

Veronatus 32, 117: tricostellus 32, 141, 172

Xenocnema spinipes 167, 183

Xenoscelinae 152
Xylochus 61
Xylophilidae 64
Xylophilus 64: nitidus 64, 160, 180
Xylotoles costipennis 65, 161, 180

Zearagytodes 26: maculifer 26

Zecillenus: alacris 21, 129, 168; tillyardi 184
Zeonidicola 50: chathamensis 50, 152, 176;
dumbletoni 50
Zeopoecilus 184
Zolini 185
Zolodininae 60
Zolodinus 60
Zomedes 61
Zopheridae 59, 102, 109
Ζorion 65, 161, 180

-194-
 NOTICES NGA PANUI
This series of refereed occasional publications has Kua whakatūria tēnei rārangi pukapuka hei
been established to encourage those with expert whakahauhau ki nga tohunga whai mātauranga
knowledge to publish concise yet comprehensive kia whakātu i nga mea e pā ana ki nga kararehe o
accounts of elements in the New Zealand fauna. ' Niu Tireni. He āhua tohunga tēnei rārangi
The series is professional in its conception and pukapuka, engari, ko te hiahia kia mārama ai te
presentation, yet every effort is made to provide tuhituhi, kia mōhio ai te maria ki nga tohu o ia
resources for identification and information that ngārara, o ia ngārara, ā, kia whakāri i te mātau-
are accessible to the non-specialist. ranga e pā ana ki a rātou.
Fauna of N.Z deals with non-marine invertebrates Κo ēnei pukapuka Fauna of New Zealand kāore e
only, since the vertebrates are well documented, pā ana ki nga kararehe moana, arā, ki nga ika, ki
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Contributions are invited from any person with
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for study. maik
āakountpat.eEwāe ohikohi-
nga kararehe e kīa ana ko te New Zealand Arthro-
Contributors should discuss their intentions with a
pod Collection hei mātakitaki māu.
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ing work; all necessary guidance will be given. tohutohu o Fauna e tika ana, kite Etita rānei, i mua
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Subscribers should address inquiries to Fauna of

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Back issues of all numbers are available, and new E toe ana nga pukapuka o mua. Mehemea e hiahia
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ten percent.

—195—
 172° 173° 174° 175° 176° 177° 178°
172° 173° 174° 175° 176° 177° 178°

35°
35°

36°
36"
πι _ ι.
ι,.,.. Ii 37°
'rιuιιι^
37° vιr,'
t uπι•
North Island ιιιιιιι-
•ιιιιιιιΙ, Ι^ιι
AK -Auckland 38°

Ι^ Ιιπιιπιιιιιιιιπιιι
38°
ΒP - Bay of Plenty ιuπιuιιιιιι πιπ_ι
CL -Coromandel ίuιιιιιιιιιιιιιππι'
ιιι ιιιιιιιιιιιιιιιιι
GB - Gisborne
ιιιιιιιιιιπιιιιιιισ 39°
ιιιιιιιιιιιιιιι
39° ιιΛιΙΙιιι ΙΙΙΙΙΙΙΙΗ , _
HB - Hawkes Bay
ND - Northland HB
Ο
ιιιιιιιιιιιιιιιιιι,ι
ΙΙΙιιιιιι ιu_
ΝΙΒΙΙΙuιn πι

9
RI - Rangitikei πιιιιιπιv, 40°
ππιιιιιιι
40° ιιιιιιιιι►
TK -Taranaki
11111111Π
Ι
TO -Taupo ιιιιιιι
Ε
OFFSHOR E ιιιιιιΡ
WA - Wairarapa ιιιιιιιι 41°
WI -Wanganui
WA ISLANDS ιι
Kermadecs ❑
^^^iίί
WN - Wellington
WO - Waikato
Ι 42°
Three Kings ❑ 42°

11W'
Chathams 40°
40° ❑

South Island
Snares ❑

BR - Buller
SD 41° Bounty ι 41°
CO - Central Otago
❑
ιτ'rr-'
'ππ
Λι/Λ.. Λrr-
DN -Dunedin Antipodes ❑ ΙΛιιιι ιΙ Οι[
FD - Fiordland
aιιππuιπι,
Aucklands ❑

KA - Kaikoura
MB - Marlborough
Campbell ❑ ΑΙΙΙΜΙΛιιιΛ ^
ΑΙΙΙΒ ΙΙΙΙΙΙι
ι 42°

MC - Mid Canterbury
0,-420 r dΙΙΙΙΙΙΙΙΙΜΒ
MK -Mackenzie _ΙΙΙΙΙΙΙΙΙΙΙΙΙι!
^ι.ιιιιι·ιι.·ιΡ . 43°
43°
ιΙ
ΙΙΙ ιΛ ι.ιι·Λιι
W .ι ϋιί ίπ ί ιι ww, ι
44° ^ΙΙΙΙΙιιΛ.ι ιιιΙιιι"
W
^1ΙΜΙΙ.··ΙΙ ···ΙΙΙι
λιΙ.ΙΙΙ ιΛΛιι Ι,
:i•iiiiinaiiiΛiiiii
·ι
ιίιιιιιπίιιuιππΜι^ι 44°

Αιιιιιιιιιιιιι•ιιιιιιΙ
a ιιιιιιιιιιιιιιιιΛιιιι 45°
45° ι.]ιiιιιιιιιιιιιΛιιιιιΙ , '
: '7ΛΛnuΛιιΙΙΙΙΙριιιΓ
■4dΙΙιΙΙιΙΙΙΙΙΙιιΙΙΙΙ=ιι
NC - North Canterbury Λ
ΩιιθιιιιιΛιιιΙιιι ιΙ
ϊ.ϋΛιι•ιιιι ιιιιιιιιι-
''ΩιιιΙΜΙΙΙι ΙΙΙΙ11Ρ
ΝΝ -Nelson 46° λι.ιιnιιι.ιιιιιιιιιιι 46°
OL -Otago Lakes
SC - South Canterbury
SD - Marlborough Sounds ^i ι• ii ^
47° 47°
SI - Stewart Island
SL -Southland
WD - Westland

167° 168° 169° 170° 171° 172° 173° 174° 167° 168° 169° 170° 171° 172° 173° 174°

Area codes and boundaries used to categorise Base-map for plottlng collection localities; this
specimen locality data (after Crosby et al. 1976) may be photocopied without copyright release

—196—
 Norfolk
lsland
30°S Kermadec
Islands

• Lord Howe
Island

SOUTH
Three Kings
lslands
ΡΑ CI FIC
35°
TASMAN
Ο C ΕΑ N
SEA NORTH
ISLAND

40°

SCALE
(km at 45°S latitude) Chatham
Islands o
0 200 400 SOUTH
45°
1 ι ι ι ι

ISLAND

' Stewart
Bounty
The Snares • Island Islands

Antipodes ,
50° lslands
Auckland
lslands

Campbell
Island THE NEW ZEALAND SUBREGlON
(excludes Lord Howe, Norfolk, and Macquarie
islands except in the context of extralimital
zoogeography)
Macquarie
pIsland
55°S
160°E 165° 170° 175° 180° 175°W

—197—
 TITLES IN PRINT / PUNA TAITARA TAA
1 Terebrantia (Insecta: Thysanoptera) • Laurence A. Mound & Annette K. Walker
ISBN 0-477-06687-9 • 23 Dec 1982 • 120 pp. $29.95
2 Osoriinae (Insecta: Coleoptera: Staphylinidae) • H. Pauline McCall
ISBN 0-477-06688-7 • 23 Dec 1982 • 96 pp. $18.60
3 Anthribidae (Insecta: Coleoptera) • B.A. Holloway
ISBN 0-477-06703-4 • 23 Dec 1982 • 272 pp. $41.00
4 Eriophyoidea except Eriophyinae (Arachnida: Acari) • D.C.M. Manson
ISBN 0-477-06745-X • 12 Nov 1984 • 144 pρ. $29.95
5 Eriophyinae (Arachnida: Acari: Eriophyoidea) • D.C.M. Manson
ISBN 0-477-06746-8 • 14 Nov 1984 • 128 pp. $29.95
6 Hydraenidae (Insecta: Coleoptera) • R.G. Ordish
ISBN 0-477-06747-6 • 12 Nov 1984 • 64 pp. $18.60
7 Cryptostigmata (Arachnida: Acari) — a concise review • M. Luxton
ISBN 0-477-06762-X • 8 Dec 1985 • 112 pp. $29.95
8 Calliphoridae (Insecta: Diptera) • James P. Dear
ISBN 0 477 06764 6 • 24 Feb 1986 • 88 pp.
- - - $18.60
9 Protura (Insecta) • S.L. Tuxen
ISBN 0 477 06765 4 • 24 Feb 1986 • 52 pp.
- - - $18.60
10 Tubulifera (Insecta: Thysanoptera) • Laurence A. Mound & Annette K. Walker
ISBN 0 477 06784 0 • 22 Sep 1986 • 144 pp.
- - - $34.65
11 Pseudococcidae (Insecta: Hemiptera) • J.M. Cox
ISBN 0 477 06791 3 • 7 Apr 1987 • 232 pp.
- - - $49.95
12 Pompilidae (Insecta: Hymenoptera) • A.C. Harris
ISBN 0-477-02501-3 • 13 Nov 1987 • 160 pρ. $39.95
13 Encyrtldae (Insecta: Hymenoptera) • J.S. Noyes
ISBN 0-477-02517-X • 9 May 1988 • 192 pp. $44.95
14 Lepidoptera — annotated catalogue, and keys to family-group taxa
J. S. Dugdale • ISBN 0-477-02518-8 • 23 Sep 1988 • 264 pp. $49.95
15 Ambositrinae (Insecta: Hymenoptera: Diapriidae) • I. D. Naumann
ISBN 0-477-02535-8 • 30 Dec 1988 • 168 pp. $39.95
16 Nepticulidae (Insecta: Lepidoptera) • Hans Donner & Chrlstopher Wilkinson
ISBN 0 477 02538 2 • 28 Apr 1989 • 92 pp.
- - - $22.95
17 Mymaridae (Insecta: Hymenoptera) • J.S. Noyes & E.W Valentine
ISBN 0-477-02542-0 • 28 Apr 1989 • 100 pp. $24.95
18 Chalcidoidea (Insecta: Hymenoptera) — introduction, and review of smaller families
J.S. Noyes & Ε.W. Valentine • ISBN 0-477-02545-5 • 2 Aug 1989 • 96 pp $24.95

—198—
19 Mantodea (Insecta), with a review of aspects of functional morphology
and biology • G.W. Ramsay • ISBN 0-477-02581-1 • 13 Jun 1990 • 96 pp $24.95
20 Bibionidae (Insecta: Diptera) • Roy A. Harrison
ISBN 0-477-02595-1 • 13 Nov 1990 • 28 pp. $14.95
21 Margarodidae (Insecta: Hemiptera) • C.F. Morales
ISBN 0-477-02607-9 • 27 May 1991 • 124 pp. $34.95
22 Notonemouridae (Insecta: Plecoptera) • I.D. McLellan
ISBN 0-477-02518-8 • 27 May 1991 • 64 pp. $24.95
23 Sciapodinae, Medeterinae (Insecta: Diptera) with a generic review of the
Dolichopodidae • D.J. Bickel • ISBN 0-477-02627-3 • 13 Jan 1992 • 74 ρρ $27.95
24 Therevidae (Insecta: Diptera) • L. Lyneborg
ISBN 0-477-02632-X • 4 Mar 1992 • 140 pp $34.95
25 Cercopidae (Insecta: Homoptera) • K.G.A. Hamilton & C.F. Morales
ISBN 0 477 02636 2 • 25 May 1992 • 40 pp. $17.95
- - -

26 Tenebrionidae (Insecta: Coleoptera): catalogue of types and keys to taxa

J.C. Watt • ISBN 0-477-02639-7 • 13 Jul 1992 • 70 pp. $27.95
27 Antarctoperlinae (Insecta: Plecoptera) • I. D. McLellan
ISBN 0 477 01644 8 • 18 Feb 1993 • 70 pp. $27.95
- - -

28 Larvae of Curculionoidea (Insecta: Coleoptera): a systematic overview

Brenda M. May • ISBN 0-478-04505-0 • 14 Jun 1993 • 226 pp. $55.00
29 Cryptorhynchlnae (Insecta: Coleoptera: Curculionidae)
C.H.C. Lyal • ISBN 0 478 04518 2 • 2 Dec 1993 • 308 pp.
- - - $65.00
30 Hepialidae (Insecta: Lepidoptera) • J.S. Dugdale
ISBN 0 478 04524 7 • 1 Mar 1994 • 164 pp.
- - - $42.50
31 Talitridae (Crustacea: Amphipods) • K.W. Duncan
lSBN 0-478-04533-6 • 7 Oct 1994 • 128 pp. $36.00
32 Sphecidae (Insecta: Hymenoptera) • A.C. Harris
ISBN 0-478-04534-4 • 7 Oct 1994 • 112 pp. $33.50
33 Moranilini (Insecta: Hymenoptera) • J.A. Berry
ISBN 0-478-04538-7 • 8 May 1995 • 82 pp. $29.95
34 Anthicidae (Insecta: Coleoptera) • F.G. Werner& D.S. Chandler
ISBN 0 478 04547 6 • 21 Jun 1995 • 64 pp. $26.50
- - -

35 Cydnidae, Acanthosomatidae, and Pentatomidae (Insecta: Heteroptera):

systematics, geographical distribution, and bioecology • M. C. Larivière
-

ISBN 0-478-09301-2 • 23 Nov 1995 • 112 pp. $42.50

36 Leptophlebiidae (Insecta: Ephemeroptera) • D.R. Towns & W.L. Peters
ISBN 0-478-09303-9 • 19 Aug 1996 • 144 pp. $39.50
37 Coleoptera: family-group review and keys to identification • J. Klimaszewski
& J.C. Watt • ISBN 0-478-09312-8 • Jul 1997 • 199 pp. $49.50

—199—
 Fauna of
New Zealand
Ko te Aitanga Pepeke
o Aotearoa

Number 37

14.

roue review
and

J. Klimaszewski & J.C. Watt

with illustrations by D.W. Helmore
POPULAR SUMMARY

CHECKLIST OF TAXA

Ν / .

REVIEW OF FAMILIES

KEYS TO TAXA

^j
\ \\. i
/

TAXONOMIC INDEX
This is a PDF facsimile of the printed publication, and is fully searchable. It is
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No ρart of this work covered by copyright may be reproduced or copied in any form
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Fauna of New Zealand website copy 2010, fnz.LandcareResearch.co.nz

Klimaszewski, J.; Watt, J.C. 1997: Coleoptera: family-group review and keys to
identification. Fauna of New Zealand 37, 199 pp.

Date of publication: 13 August 1997

Fauna of New Zealand, ISSN 0111-5383; 37

ISBN 0-478-09312-8

New Zealand Coleoptera. Scanned images provided by Stephen Pawson

(www.bugz.org.nz). OCRed text corrected for this searchable PDF by Trevor Crosby,
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