BIOTAXONOMY OF TEPHRITOIDEA
Isr. J. Entomol. Vol. 35-36, 2005/6, pp. 163-196
Catalog and Revised Classification of the Gastrozonini
(Diptera: Tephritidae: Dacinae)
Damir Kovac,1 Patrick Dohm,1 Amnon Freidberg2 and
Allen L. Norrbom3
ABSTRACT
A catalog of Gastrozonini is presented based on the recent world catalog of
Tephritidae and subsequent taxonomic and nomenclatural changes. The
Gastrozonini are defined taxonomically and biologically, primarily as the
Poaceae-infesting Tephritidae other than Acanthonevrini. Although
morphological synapomorphies are scant, the group is probably a
monophyletic unit based primarily on host plant records and a few other
potential synapomorphies such as characters of the antenna, wing and
terminalia. Larvae of the gastrozonine species that we studied have a ridge on
the ventral tubercle of the caudal segment, confirming the relationship of
Gastrozonini with Dacini and Ceratitidini. A key to all included genera of
Gastrozonini is provided, and new taxonomic and nomenclatural changes
introduced in the catalog are explained. The current classification comprises 27
1
Forschungsinstitut Senckenberg, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany. E-mail
(Kovac): Damir.Kovac@senckenberg.de; E-mail (Dohm): patrick.dohm@senckenberg.de. 2Department
of Zoology, Faculty of Life Sciences, Tel Aviv University, Tel Aviv 69978, Israel. E-mail:
afdipter@post.tau.ac.il. 3Systematic Entomology Laboratory, USDA, c/o National Museum of Natural
History, MRC 168, Washington, DC 20013-7012, USA. E-mail: anorrbom@sel.barc.usda.gov.
BIOTAXONOMY OF TEPHRITOIDEA
genera (of which about half have confirmed host records) and 137 species, 14
of which are Afrotropical and probably comprise a monophyletic subgroup.
The remaining group of genera extends from the southeastern part of the
Palaearctic Region through the entire Oriental Region into the Australasian
Region, as far south as Australia, but is predominantly Oriental. The following
new combinations are made: Acroceratitis aberrata Hardy, Acroceratitis
adnata Hardy, Chelyophora maai Chen, Ceratitoides namtamai Hancock, and
Ceratitoides sikhimensis Hancock are transferred from Ceratitoides to
Pardalaspinus, and Carpophthoromyia cinereofasciata Meijere is transferred
from Ceratitoides to Proanoplomus. Acrotaeniostola longicauda Wang,
previously synonymized under Acrotaeniostola morosa Hering, is reinstated
from synonymy.
INTRODUCTION
The motivation to prepare this publication comes from our recently growing interest in the
Poaceae (Gramineae)-breeding fruit flies (Tephritidae: Dacinae: Gastrozonini), often called
grass-breeding or bamboo-shoot fruit flies (Wang, 1998; Hancock, 1999; Hancock and Drew,
1999). Although confirmed published host records are available for a relatively small number of
species (Allwood et al., 1999), all the species of this group probably breed in Poaceae. In the
Oriental Region, where the great majority of the species occurs, existing host records indicate
that most or all of the species are bamboo breeders.
Almost nothing is known about the biology of the Gastrozonini, apart from the confirmed
and suspected host records (Hancock and Drew, 1999) and an isolated description of the
behavior of one species, Cyrtostola limbata (Hendel) (Kovac and Azarae, 1994). The bulk of
the published information on the group is taxonomic, but the taxonomy of the group is still in
flux, and even the taxonomic limits of the group are uncertain. For example, Hancock (1999),
when discussing the relatively few Afrotropical taxa together with some of the taxa from the
Oriental Region [an artificial combined group] noted: “The resemblance of Bistrispinaria
[African (our addition)] to the Anoplomus complex [‘Oriental’ (our addition)] suggests that all
may breed in grasses and thus all belong in tribe Gastrozonini”. However, he has not applied
this conclusion and, in two papers that appeared in the same issue, retained the Anoplomus
group (including the Afrotropical taxa) in the Ceratitidini (Hancock, 1999), separate from the
Gastrozonini (Hancock and Drew, 1999).
As with many other groups of Tephritidae, the lack of comprehensive phylogenetic analysis
and apparent homoplasy in conventional taxonomic characters obscures the true taxonomic
relationships of this group. Much more research is still required, especially biological and
molecular, as well as a comprehensive study of the male and female terminalia, before a
satisfactory classification can be achieved. Nevertheless, the information gathered recently,
especially in the publications cited above and also through the unpublished results of our work
in Asia and Africa, indicates that the group treated in the present publication is most probably
monophyletic.
Gastrozonini are not the only fruit fly group breeding in Poaceae. A group of genera
referable to the Acanthonevrini and centered around Acanthonevra Macquart, Ptilona Wulp
and Felderimyia Hendel, is also associated with bamboo in the Oriental Region and adjacent
areas of the Palaearctic and Australasian Regions, although their association is probably
primarily with dead or decaying bamboo. The taxonomic and phylogenetic relationships of the
Acanthonevrini and Gastrozonini are not fully understood, but they are currently classified in
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different subfamilies and do not appear to be closely related (Korneyev, 1999). Moreover, there
is no evidence supporting an assumption that the bamboo breeding Acanthonevrini form a
monophyletic group, and their relationships with the rest of the Acanthonevrini (some of which
are known to have other host plant associations) are not clear.
Korneyev (1999) included the Gastrozonini, Ceratitidini and Dacini in the subfamily
Dacinae. Norrbom et al. (1999) gave these groups a lower rank: Gastrozonina, Ceratitidina and
Dacina, together comprising the tribe Dacini. Here, we prefer to follow Korneyev (1999)
because his classification was based on a cladistic analysis. Whatever the nominal rank of these
more or less coherent groups is, the relationships among them are still unresolved.
The monophyly of the subfamily Dacinae is supported by morphological and biochemical
data, although some characters have not been studied thoroughly in the Gastrozonini
(Korneyev, 1999). Hancock (1986) and Foote et al. (1993) suggested that the Dacinae are
monophyletic based on the shape of the posterodistal lobe of cell bcu (often narrower at base
than medially), spermathecae number reduced to two (considered a synapomorphy of Dacini +
Tephritinae by Hancock, 1986), and surstyli shape. Korneyev (1999) also considered the large
proctiger of the male to be a synapomorphy of the subfamily. Species of Dacini and Ceratitidini
whose larvae have been described (no larvae of Gastrozonini have been described yet) have a
ridge across the large tubercle ventral to the posterior spiracle on the caudal segment, an
apomorphic state not reported in any other Tephritidae (Carroll, 1992). Our unpublished data on
ten species of Gastrozonini indicate that this character is also common to this tribe, confirming
its relationship to the Ceratitidini and Dacini. Kitto (1983) suggested the close relationship of
the few Dacini and Ceratitidini included in his immunological study (no Gastrozonini were
studied), and the four species (2 Dacini, 1 Ceratitidini, 1 Gastrozonini) included by Han and
McPheron (1997) were grouped together in their neighbor-joining tree.
Relationships within the Dacinae have not been rigorously analyzed. The Dacini and
Ceratitidini each appear to be monophyletic, but the Gastrozonini were previously considered
to possibly be paraphyletic (Norrbom et al., 1999; Korneyev, 1999). We now consider the group
to be monophyletic (see diagnosis), although some adjustments (e.g., the inclusion of additional
taxa) may still be necessary.
All this means that diagnosing the Gastrozonini is difficult, although we have nevertheless
attempted to do so in the short diagnosis of the group that follows. The best diagnostic character
of the Gastrozonini is their association with Poaceae. Combining the confirmed published
records (Hancock and Drew, 1999) with our unpublished records has resulted in a host list for
13 of the 27 genera (see below). Most Gastrozonini genera, as presently conceived, appear to be
homogenous units, and the host consistency observed to date within the genera for which host
records are available for multiple species suggests that host plant association is constant within
genera. Several apomorphic or potentially apomorphic morphological characters (some are of
uncertain polarity) also support the monophyly of the group, although not all are consistently
present. These include: 1st flagellomere often with distinctly raised dorsoapical point; arista
almost always plumose; setulae on pedicel long, dense and dark; wing pattern almost always
banded, usually more or less Ceratitis-like; male proctiger enlarged, covering entire epandrium
in ventral view; and aculeus usually very broad, flattened, strongly sclerotized and pointed or
relatively elongate, narrow and pointed.
All gastrozonine genera conform to the diagnosis below. Those without host records are
either similar to genera with host records or, at least, exhibit some of the crucial diagnostic
traits, such as the shape of the antenna and aculeus. As stated earlier, most of these genera, even
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BIOTAXONOMY OF TEPHRITOIDEA
if not definitively included in the Gastrozonini by Hancock (1999), were implied by him to
belong there. The only debate concerns the wasp mimic, Ichneumonopsis Hardy, which was
included by Hancock (1986) and Drew and Hancock (1994) in the Dacina (a large taxon of
wasp-like flies). In addition to this shared mimicry (Freidberg, unpublished observations), this
genus also shares with the Dacina reduced chaetotaxy, but it does not share with this group
other, more important, synapomorphies, such as those of the wing venation (Radhakrishnan,
2000) and preabdomen, and therefore we continue to classify it in the Gastrozonini. Reduced
chaetotaxy has occurred multiple times in tephritid evolution, including other wasp mimics,
such as the Adramini, Toxotrypana Gerstaecker, and some Pseudophorellia Lima (Norrbom,
2006, this volume).
Gastrozonini that breed in bamboo are attracted to damaged but living bamboo plants and
probably develop, at least partly, in live tissues. As such they should be considered as actual or
potential pests of bamboo, a commodity of great importance in southeast Asia. As their biology
is just beginning to be carefully explored, further study is necessary before the damage they
inflict to bamboo, or other Poaceae, can be assessed. Many species have been recorded to be
attracted to freshly cut bamboo (Hancock and Drew, 1999), and this behavior has been
suggested as diagnostic for Gastrozonini. Our own observations generally support this notion.
However, we would urge scientists to be cautious and use this behavioral trait only as an
indication to possible taxonomic assignments of Tephritidae attracted to bamboo, since we have
observed other tephritids, such as the distantly related Adrama Walker, attracted to freshly cut
bamboo.
The Gastrozonini are primarily tropical, with the largest number of species in the Oriental
Region and a few in the Afrotropical, southeastern Palaearctic and Australasian Regions. The
Palaearctic and Australasian species show close affinities to the Oriental fauna, whereas the
four Afrotropical genera are very similar to each other and may comprise a separate
monophyletic subgroup. However, the latter hypothesis should be tested.
The purpose of this publication is to compile in one source the previously scattered
nomenclatural and taxonomic information on the Gastrozonini subsequent to the world catalog
and database of the Tephritidae (Norrbom et al., 1999; Norrbom, 2004), and provide a single
key to all of the known genera. We have entered taxonomic and nomenclatural changes from
two kinds of sources: 1) recent publications not available at the time the world catalog was
produced (i.e., ranging between 1999-2004); and 2) changes proposed by us herein, that we
have kept to a minimum. Hence, almost all the many changes suggested by Hancock since 1999
(Hancock, 1999; Hancock and Drew, 1999) have been incorporated without critical checking,
and in only two cases did we introduce new changes. These latter changes are explained in a
special section following the catalog.
The format of this catalog is similar to that of the world catalog, with mostly the same
abbreviations and acronyms. Each genus entry contains a header with the name of the valid
genus in bold, and for each name (valid name, synonyms, homonyms, misspellings,
misidentifications) pertaining to that genus the following information: Original name, author,
year, page number, type-species, kind of designation, and as applicable, note information. Each
species entry contains a header with the name of the valid species in bold, and for each name
(valid name, synonyms, homonyms, misspellings, misidentifications) pertaining to that species
the following information: Original combination, author, year, page number, type data (kind of
type, sex of type, depository, type locality), and as applicable, note information. The
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Kovac et al. / Catalog and classification of Gastrozonini
distribution is also given under the valid name. A host catalog covering most Gastrozonini was
published by Hancock and Drew (1999), and it is not repeated here. The genera that have been
positively associated with Poaceae (i.e., at least one species of a genus has a confirmed rearing
record; an asterisk (*) denotes new rearing records) are Acroceratitis, Acrotaeniostola,
Bistrispinaria, *Carpophthorella, Chaetellipsis, Cyrtostola, *Enicoptera, Gastrozona,
Paragastrozona, Paraxarnuta, Phaeospilodes, Taeniostola, Xanthorrhachis, and there are
indications for such an association for several other genera.
A key to the genera of Gastrozonini is provided to help identify the genera according to the
new classification proposed here. This key essentially combines the two keys to genera of
Hancock (1999) and Hancock and Drew (1999), respectively, with necessary modifications and
adding to them Ichneumonopsis. We have tested about 65 species against the key, representing
the great majority of genera, and consequently made some minor adjustments, including style,
terminology and rearrangement. Representatives of only Chelyophora, Phaeospila and
Sinanoplomus have not been tested. During the process, we have encountered some difficulties,
primarily caused by the vague definition and limits of some genera. A particularly difficult area
of the key is the group of genera clustered around Gastrozona (couplets 13-18), and a
particularly interesting section is formed by the African taxa. Because the African taxa are
placed for the first time in context with all ‘Asian’ Gastrozonini, and because no cladistic
analysis is available for the Gastrozonini, we have refrained from introducing drastic changes to
Hancock’s (1999) key. However, the perspective we have gained during the process indicated
that a) the African taxa are very similar to each other and probably form a monophyletic group;
b) this group may not necessarily be the sister group of all the Oriental Gastrozonini but a
subgroup thereof; and c) the concepts of the four currently recognized African genera are not
significantly different, and all African species should probably be united under one genus
(Bistrispinaria), which nevertheless lacks good diagnostic characters at present. This would
better be done as part of a future phylogenetic analysis of the Gastrozonini. Hancock’s keys to
species (loc. cit.) generally work well, although for better results in certain genera (e.g.,
Acrotaeniostola), Wang’s (1998) keys should also be consulted.
Terminology follows McAlpine (1981) and White et al. (1999). In the wing pattern we used
the term transverse for bands that are oriented more or less perpendicular to the longitudinal
axis of the wing, longitudinal for bands parallel to this axis, and oblique for transverse bands
that are diagonal to this axis.
List of abbreviations used in the catalog (slightly modified from Thompson, 1999)
A
—
adult
B
—
both sexes
E., e.
—
east, eastern
Emend.
—
emendation
F
—
female
ft.
—
feet
HT
—
holotype
I., Is.
—
island, islands
Incosp.
—
incorrect original spelling
LT
—
lectotype
M
—
male
m
—
meter(s)
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BIOTAXONOMY OF TEPHRITOIDEA
mi.
Misid.
Missp.
MO
Mt., Mts.
N. N.
N., n.
N. Comb.
NE., ne.
nr.
NW., nw.
OD
R.
S., s.
SE., se.
ssp.
ST
T
W., w.
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
mile(s)
misidentification
misspelling
monotypy
mount or mountain(s)
new name
north, northern
new combination
northeast, northeastern
near
northwest, northwestern
original designation
river
south, southern
southeast, southeastern
subspecies
syntype
type
west, western
List of collections housing type material and their acronyms
Baker — C.F. Baker, personal collection. (The relevant specimens are actually in MCSNM)
BAUC — Beijing Agricultural University, Beijing
BBM — Bernice P. Bishop Museum, Honolulu
BMNH — The Natural History Museum, London
CAS — California Academy of Sciences, San Francisco
DEI — Deutsches Entomologisches Institut, Müncheberg, Germany (formerly: Eberswalde)
HUS — Entomological Institute, Hokkaido University, Sapporo, Japan
IZAS — Institute of Zoology, Academia Sinica, Beijing
KUB — Kasetsart University, Bangkok, Thailand
MCSNG — Museo Civico di Storia Naturale, Genoa, Italy
MCSNM — Museo Civico di Storia Naturale, Milan
MNHNP — Museum National d’Histoire Naturelle, Paris
MNM — Magyar Termeszettudomanyi Muzeum Allattara, (Hungarian Natural History
Museum), Budapest
MRAC — Musée Royal de l’Afrique Centrale, Tervuren, Belgium
NMBZ — Natural History Museum, Bulawayo, Zimbabwe
NMW — Naturhistorisches Museum Wien
NRS — Naturhistoriska Riksmuseet, Stockholm
NSWA — New South Wales Agricultural Scientific Collection Trust, Rydalmere, NSW,
Australia
NTU — National Taiwan University, Taipei
PAN — Polish Academy of Science, Museum of the Institute of Zoology, Warsaw
PUCP — Punjab University, Chandigarh, Punjab, India
RNH — Nationaal Natuurhistorische Museum, Leiden, Netherlands
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SANC — South African National Collection of Insects, Pretoria
UOPJ — Entomological Laboratory, University of Osaka, Osaka, Japan
USNM — United States National Museum of Natural History, Washington, DC.
UZMH — Zoological Museum, Finnish Museum of Natural History, University of Helsinki,
Helsinki
ZFMK — Zoologisches Forschungsinstitut und Museum “Alexander Koenig”, Bonn, Germany
ZMAN — Zoologisch Museum, Universiteit van Amsterdam, Amsterdam
ZMHU — Museum für Naturkunde der Humboldt Universität zu Berlin, Berlin
ZSI — Zoological Survey of India, National Zoological Collection, Calcutta, India
ZSZMH — Zoologisches Staatsinstitut und Zoologisches Museum, Hamburg, Germany
Additional material that was used by us for the preparation of this publication is deposited in the
Senckenberg Museum, Frankfurt/Main, Germany (SMF) and Zoological Museum, Tel Aviv
University, Tel Aviv, Israel (TAUI).
DIAGNOSIS OF GASTROZONINI
No single feature can distinguish all Gastrozonini. Hence this detailed diagnosis (aranged in the
format of a description) contains both plesiomorphic and apomorphic characters (marked by a
(= apomorphy) or p (= plesiomorphy) where their polarity appears obvious). Of these, the most
important appears to be the association with Poaceae, and more specifically with bamboo
(Asian species) or non-bamboo hosts (African species). It should be noted that character
polarities were tentatively assigned based on comparison with other Dacinae and other
Tephritidae. The important morphological characters are as follows.
Head. Antenna: 1st flagellomere often with distinctly raised dorsoapical point (a), or the
meeting point of the straight dorsal margin with the rounded apical edge sharp, or 1st
flagellomere tapered, ending in distal point. In ten genera (Carpophthorella, Chaetellipsis,
Cyrtostola, Dietheria, Enicoptera, Gastrozona, Ichneumonopsis, Paragastrozona,
Pardalaspinus, Proanoplomus) 1st flagellomere more or less rounded distally. Arista never
bare, almost always plumose (polarity uncertain), with long rays, and plumosity usually
exceeding or equal to antennal width. Only rarely rays short, and arista pectinate
(Chaetellipsis), or pubescent (Paragastrozona). Setulae on pedicel long, dense and dark in all
African species and in many, but not all Asian species (a?). Chaetotaxy: Full complement of
setae usually present (p), and setae usually “strong” (long and thick) and dark. Setae on anterior
part of gena occasionally strongly developed as major setae (e.g., Acrotaeniostola). Some major
setae, e.g., ocellar or frontal setae, sometimes reduced or lacking.
Thorax. Scutellum usually flat, sometimes convex, always setulose. Color pattern, particularly
on mesonotum, often composed of contrasting pale and dark areas, obscured or not by areas of
microtrichia and pale or dark setulae. Chaetotaxy: Usually full complement of “strong” setae
present (p), but some major setae occasionally lacking. The usual and maximum number of
scutellar setae is 2, but in three genera only 1 is present.
Legs. No overt features.
Wing. Venation: Usually similar to Ceratitidini and Dacini in relatively long and basally
constricted posterodistal lobe of cell bcu (although lobe shorter than in Dacini), but different
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BIOTAXONOMY OF TEPHRITOIDEA
from most Dacini in narrow cell bm (p). Venation rarely of unusual form (Enicoptera - a).
Pattern: Almost always banded, usually without, sometimes with Ceratitis-like black subbasal
streaks, with various subtypes exhibiting various combinations of transverse, longitudinal, and
oblique bands. Pattern sometimes reduced to spots that seem to conform to the abovementioned subtypes. Wing pattern types of rare occurrence (a) are: merely longitudinal bands,
including dimidiate pattern (e.g., Xanthorrachis, Ichneumonopsis); reticulate or reticulatebanded (e.g., Phaeospilodes); and the complicated and colorful pattern of males of some
Chaetellipsis species.
Abdomen. Often with distinct pattern of microtrichia, primarily on tergites 3 and 5. Male
tergite 2 often greatly enlarged, covering lateral margin of tergite 3 and with large specialized
areas laterally (a). Male proctiger enlarged, covering entire epandrium in ventral view (a).
Female with two spermathecae (p; possible synapomorphy of Dacinae). Female aculeus of
various shapes (a). In Gastrozona, Chaetellipsis, and a few other genera the aculeus is relatively
elongate and narrow, pointed apically, and often with preapical steps and occasionally serration
(polarity uncertain). In Acrotaeniostola, Acroceratitis, and many other genera, the aculeus is
relatively broad and flattened, usually strongly sclerotized and pointed, but occasionally
rounded apically (a; this is the dominant shape in Gastrozonini, and it is relatively little varied).
In Dietheria the tip of the aculeus is bifid (a).
KEY TO THE GENERA OF GASTROZONINI
1. Extremely wasp-like species, with petiolate abdomen, spinose forefemur and dimidiate
wing pattern (anterior half of wing yellow, remaining part hyaline), chaetotaxy reduced: one
setula-like frontal seta, one small orbital seta and 1 scutellar seta present; ocellar,
dorsocentral, acrostichal, intra-alar, postpronotal and pleural setae, except one anepisternal
seta, lacking ................................................................................................ Ichneumonopsis
–. Not extremely wasp-like species, abdomen not petiolate, forefemur not spinose, wing with
different pattern, chaetotaxy usually not reduced, if reduced, then only one or two of abovementioned setae lacking ...................................................................................................... 2
2. Vein R2+3 bowed anteriorly towards vein R1 near apex, meeting vein R1 or, sometimes, split
into vein R2 and vein R3, thus forming small, peculiar cell just beyond pterostigma; vein M
sinuous, bowed anteriorly beyond apex of cell dm and almost meeting vein R4+5 at wing
apex; pterostigma elongate, ending considerably beyond mid-length of wing and opposite
crossvein DM-Cu; crossvein DM-Cu strongly oblique anterodistally ................ Enicoptera
–. Veins R2+3 and M not sinuous; pterostigma not elongate ending at about mid-length of wing
and approximately opposite crossvein R-M; crossvein DM-Cu not strongly oblique
anterodistally ......................................................................................................................... 3
3. Head with 4–16 pairs of frontal setae or, in some males, frontal and orbital setae absent; wing
without oblique brown band from vein A1+Cu2 across crossvein R-M to costa; anepisternum
with whitish band across dorsal part from postpronotal lobe to wing base; sexes often
strongly dimorphic in wing pattern or color ......................................................................... 4
–. Head usually with 1-3 pairs of frontal setae, if with 4-5 (pairs) then wing with partial or
complete oblique brown band from vein A1+Cu2 across crossvein R-M to costa; if this band
reduced to isolated spots then anepisternum almost entirely yellowish, sexes at most weakly
dimorphic in wing pattern or color ....................................................................................... 5
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Kovac et al. / Catalog and classification of Gastrozonini
4. Male with frontal and orbital setae present; female with oblique brown band in posterior part
of wing, from vein A1+Cu2 across distal part of cell dm reaching or almost reaching
subapical band ............................................................................................. Carpophthorella
–. Male with frontal and orbital setae absent; female without oblique brown band from vein
A1+Cu2 to subapical band, but with isolated band over crossvein DM-Cu to posterior wing
margin....................................................................................................................Chaetellipsis
5. Wing with 2 or 3 yellow longitudinal bands, along costa, vein M and usually vein Cu1;
ocellar setae usually vestigial; male tergite 2 enlarged, with large unusually textured area
laterally (secretional? stridulatory?), covered only by dense and erect minute setulae, unlike
other abdominal setulae ........................................................................................................ 6
–. Wing pattern different, dark with hyaline spots and indentations or with transverse or
oblique yellow to brown bands, or reduced to 1-2 transverse bands plus more or less isolated
small spots; ocellar setae usually well-developed; male tergite 2 enlarged or not, with or
without such specialized area ............................................................................................... 7
6. Head nearly quadrate, with frons almost horizontal; body less robust, with male abdomen
about 1.5 times as long as wide ........................................................................... Galbifascia
–. Head distinctly higher than long, with frons sloping; body more robust, with male abdomen
about as long as wide ...................................................................................... Xanthorrachis
7. Scutum, scutellum and abdomen entirely and densely covered by short black setulae, usually
strongly contrasted with paler background (some scutal setulae occasionally appearing pale
in certain viewing angles) and with about 15-20 irregular rows of setulae between acrostichal
setae; ventral margin of scutellum extensively setulose; wing pattern reduced, comprising
broad and complete, nearly transverse band over pterostigma and crossvein R-M, and more
or less complete, converging band over crossvein DM-Cu, as well as few small spots and
occasionally yellowish base of wing ................................................................. Paraxarnuta
–. Scutum, scutellum and abdomen not so densely black setulose, setulae usually not strongly
contrasted with background, usually pale, if black then no more than 12 rows of setulae
between acrostichal setae, and wing pattern not reduced .................................................... 8
8. Wing pattern comprising large preapical brown area posterodistally (usually covering both
crossveins R-M and DM-Cu, sometimes only DM-Cu), with long costal band over
pterostigma and most or all cell r1 and occasionally with short band along central part of cell
dm, both attached to main dark area, and with narrow dark crescentic apical spot partly
(male) or entirely (female) separated from main dark area by crescentic hyaline gap;
crossvein DM-Cu anterodistally oblique; lacking ocellar, postocellar, postpronotal,
presutural and dorsocentral setae .................................................................... Rhaibophleps
–. Wing pattern and venation different, wing without such longitudinal bands; chaetotaxy
more complete ..................................................................................................................... 9
9. Wing banded, with all bands generally parallel and oblique, except sometimes (in
Taeniostola) on apical third of wing, where 2-3 bands may converge ............................. 10
–. Wing banded or not, if banded then bands on apical half to two-thirds of wing either
converging or generally parallel and transverse, usually including four bands arranged more
or less in the form of a small “v” within a large “v”; wing pattern sometimes reticulate or
reticulate-banded ............................................................................................................... 12
10. Chaetotaxy: 1 scutellar seta, 1 frontal seta, 1 orbital seta ..................................... Dietheria
–. Chaetotaxy: 2 scutellar setae, 2-5 frontal setae, 2 orbital setae ......................................... 11
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BIOTAXONOMY OF TEPHRITOIDEA
11. First flagellomere distinctly pointed dorsoapically; midtibia with 2 subequal apical spines;
postpronotal lobe and scutum with isolated black spots .................................. Spilocosmia
–. First flagellomere indistinctly pointed or rounded apically; midtibia with one apical spine;
postpronotal lobe and scutum with or without black vittae, but without black spots ...........
............................................................................................................................ Taeniostola
12. African; probably not associated with bamboo ................................................................. 13
–. Asian; known or suspected to be associated with bamboo ................................................ 16
13. Wing with 2-5 long costal spines at end of vein Sc and distinct dark spots and streaks at
base, including elongate streak in cell c; scutellum mostly black with distinct longitudinal
medial yellow spot at base, pair of basolateral yellow lines and usually pair of subapical
yellow lines ..................................................................................................... Bistrispinaria
–. Wing usually with at most one long costal spine at end of vein Sc and indistinct spots and
streaks at base, usually without elongate streak in cell c; scutellum entirely yellow, yellow
to white with one or three black apical or subapical spots, or mostly or entirely black,
without distinct longitudinal medial yellow spot at base .................................................. 14
14. Wing with transverse band from cell sc, across central part of cell br and basal half of cell
dm to hind margin, united at vein M and apex of pterostigma with band along crossveins
DM-Cu and R-M ............................................................................................... Ceratitoides
–. Wing with oblique band from beyond cell sc, across crossvein R-M and middle of cell dm,
leaving distal half of pterostigma and basal part of cell r1 pale or hyaline and usually free
from band along crossvein DM-Cu; oblique band usually entirely disconnected from basal
area of pattern .................................................................................................................... 15
15. Scutum yellowish, with pair of black spots posterolaterally in both sexes; scutellum with
apical spot usually small, rounded, restricted to area between apical setae; abdomen yellow
to brown, at most with brownish bands; mediotergite yellow to red brown.................
............................................................................................................................Leucotaeniella
–. Scutum brownish black to black, if mostly yellow in male then black areas present
dorsocentrally in posterior part, scutellum with apical spot large, quadrate, usually about
half as long as scutellum, sometimes almost as large as scutellum, and abdomen with black
transverse bands; mediotergite shiny blackish brown to black .......................... Clinotaenia
16. Wing with dark claw-like marginal band, extending from level of crossvein H, where it
occupies entire wing width, covering about half of anal lobe, to wing tip, including
crossvein R-M, and with two converging and more or less well-developed subapical and
posterior apical bands .................................................................................... Sinanoplomus
–. Wing pattern different ....................................................................................................... 17
17. Wing with brown band over crossvein R-M (discal band) oblique, reaching or directed
towards costa beyond end of cell sc, separated from it by hyaline band or indentation, if
ending close to apex of cell sc then 1st flagellomere apically rounded and band across
crossvein DM-Cu connected with subapical or apical brown markings ........................... 18
–. Wing with brown or yellow band over crossvein R-M (discal band), when present,
transverse or nearly so, usually reaching or directed towards costa within cell sc, if ending in
cell r1 at apex of cell sc then 1st flagellomere dorsoapically pointed and band across
crossvein DM-Cu not joined with subapical and apical dark markings; discal band
sometimes not extending over crossvein R-M (subapical band is instead) ....................... 22
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18. Scutum with four longitudinal black vittae, middle pair aligned with acrostichal setae; head
with two frontal setae ........................................................................................... Cyrtostola
–. Scutum black, or with obscure lyre-like pattern, or almost entirely yellow, or with two or
three (rarely 0 or 5) longitudinal black vittae, lateral pair often interrupted, medial vitta
often absent, particularly in males; head with 2-5, usually 3-5, frontal setae ................... 19
19. Postpronotal seta absent; 2 frontal setae ............................................................. Anoplomus
–. Postpronotal seta present; 3-5 frontal setae ....................................................................... 20
20. Head about 1.1 times as high as long; aculeus flat, broad and non-setulose; wing with cell bc
fulvous; arista pubescent (in type species) .................................................. Paragastrozona
–. Head 1.33-1.50 times as high as long; aculeus variable but often narrow and with long
setulae preapically; cell bc variable, but usually mostly or entirely hyaline, arista usually
plumose .............................................................................................................................. 21
21. Mesonotum usually predominantly yellow or with more or less balanced pattern of yellow
and darker parts, often including black or blackish medial vitta broadened anterior to
scutellum in female and trace of such vitta in male, connected to median scutellar spot, or
transverse brownish prescutellar band; aculeus elongate and with long setulae preapically
............................................................................................................................ Gastrozona
–. Mesonotum predominantly black or blackish, without such pattern, aculeus broad .............
....................................................................................................................... Proanoplomus
22. Scutum without distinct yellow or white prescutellar markings; scutellum fulvous to yellow
or medially darkened but without posterior black patches, or scutum and scutellum
predominantly shiny black ................................................................................................ 23
–. Scutum with distinct yellow or white prescutellar markings, if not than scutellum with three
large posterior black patches or mostly black posteriorly ................................................. 25
23. Wing predominantly dark with few, relatively large, hyaline spots and indentations, not
banded.......................................................................................................................Phaeospila
–. Wing pattern banded .......................................................................................................... 24
24. Wing bands generally parallel except when narrow spiral costal band present from wing tip
to base of pterostigma, then to tip of vein A1+Cu2, sometimes connected to band over
crossvein DM-Cu ......................................................................................... Acrotaeniostola
–. Wing bands generally convergent at distal 0.66 of wing; costal band present, but neither
narrow, nor spiral ................................................................................. Pardalaspinus (part)
25. Wing pattern reticulate or reticulate-banded, brown, with at most few small indistinct
yellow brown patches, with hyaline spots and indentations at least in apical half, without
subapical or costal bands ............................................................................... Phaeospilodes
–. Wing pattern banded, yellow or yellow and brown, if bands not evident in apical half, then
pattern there diffuse ........................................................................................................... 26
26. Two postpronotal setae; 1st flagellomere rounded apically .............................. Chelyophora
–. One postpronotal seta; 1st flagellomere rounded or pointed apically ................................ 27
27. Scutum and scutellum predominantly shiny black; 1st flagellomere usually rounded apically
................................ Pardalaspinus (part, at least including the type species — laqueatus)
–. Scutum and scutellum not predominantly shiny black; 1 st flagellomere pointed
dorsoapically ..................................................................................................... Acroceratitis
173
BIOTAXONOMY OF TEPHRITOIDEA
CATALOG OF GASTROZONINI
1. ACROCERATITIS
Acroceratitis Hendel, 1913b: 82. Type species: plumosa Hendel (OD). Latest reference: Hancock and
Drew, 1999: 638.
Stictaspis Bezzi, 1913: 102. Type species: ceratitina Bezzi (OD). Synonymy: Hardy, 1973: 217.
Acroceratis Shinji, 1940a: 163, Missp. Acroceratitis Hendel.
Acroceratitis bilineata
Chelyophora bilineata Meijere, 1914: 205. HT F (ZMAN), Indonesia. Java: Semarang. Type data:
Hardy, 1988: 80. Latest reference: Hancock and Drew, 1999: 640.
Distribution: Indonesia (Java), w. Malaysia (Selangor).
Acroceratitis bimacula
Acroceratitis bimacula Hardy, 1973: 223. HT M (KUB), Thailand. Nakhon Ratchasima: Pak Chong.
Latest reference: Hancock and Drew, 1999: 642.
Distribution: India (Andaman and Nicobar Is.), China (Guangxi, Yunnan), Thailand, Laos, Vietnam.
Acroceratitis ceratitina
Stictaspis ceratitina Bezzi, 1913: 103. HT F (ZSI), India. Bihar: Parasnath, 4400 ft. Latest reference:
Hancock and Drew, 1999: 643.
Acroceratitis cognata Hardy, 1973: 225. HT M (BBM), Thailand. Tak: Mae Sot Dist., Huai Muang,
Canton, 200 m. Synonymy: Hancock and Drew, 1999: 643.
Chelyophora clavifera Hering, 1938c: 7. HT F (NRS), Burma. s. Shan: Inle Lake, s. end, Taungdo,
900 m. Synonymy: Hancock and Drew, 1999: 643.
Chelyophora gladiella Munro, 1938: 22. ST B (ZSI), India. Bihar: Pusa. Type data: Kapoor 1993: 93.
Synonymy: Hancock and Drew, 1999: 643.
Acroceratitis biseta Wang, 1998: 24. HT M (IZAS), China. Yunnan: Xishuangbanna (20ºN 100.8ºE).
Synonymy: Hancock and Drew, 1999: 774.
Distribution: India (Uttar Pradesh, Bihar, W. Bengal, Assam), China (Yunnan), Burma, Thailand.
Acroceratitis distincta
Chelyophora distincta Zia, 1964: 47. HT M (IZAS), China. Yunnan: Kin-ping [Jinping], 370 m. Latest
reference: Hancock and Drew, 1999: 645.
Phaeospilodes fritilla Hardy, 1973: 199. HT M (KUB), Thailand. Phra Nakhon: Bangkhen [Bang
Khen, 13°52'N 100°36'E]. Synonymy: Wang, 1998: 43; Hancock and Drew, 1999: 645.
Phaeospilodes distincta: Wang, 1998: 43 (redescription; China).
Distribution: ne. and s. India, China (Yunnan), Laos, Thailand, Vietnam.
Acroceratitis hardyi
Acroceratitis hardyi Hancock and Drew, 1999: 647. HT M (BMNH), Malaysia. Selangor: Ulu Langet,
nr. Kuala Lumpur.
Distribution: s. Thailand, w. Malaysia (Selangor).
Acroceratitis histrionica
Chelyophora histrionica Meijere, 1914: 205. ST B (ZMAN), Indonesia. Java: Buitenzorg [Bogor];
and Semarang. Inference of holotype by Hardy (1973: 227, 1988: 80) invalid. Latest reference: Hancock
and Drew, 1999: 649.
Distribution: Sri Lanka, Thailand, Laos, w. Malaysia, Indonesia (Java).
Acroceratitis incompleta
Acroceratitis incompleta Hardy, 1973: 227. HT M (BBM), Thailand. Chiang Mai: Doi Suthep,
1278 m. Latest reference: Hancock and Drew, 1999: 650.
Distribution: China (Yunnan), n. Thailand, Laos.
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Acroceratitis nigrifacies
Acroceratitis nigrifacies Meijere, 1924: 36. LT M (ZMAN), Indonesia. Sumatra: Barat, Padang
Plateau, Buo. Lectotype designated by inference of holotype by Hardy (1988: 81). Latest reference:
Hancock and Drew, 1999: 652.
Distribution: w. Malaysia, Indonesia (Sumatra).
Acroceratitis plumosa
Acroceratitis plumosa Hendel, 1913b: 82. ST F (DEI, NMW), Taiwan. Kankau. Type data (Hardy,
1968: 108). Latest references: Wang 1998: 25; Hancock and Drew, 1999: 653.
Distribution: China (Zhejiang, Yunnan, Hainan), Taiwan.
Acroceratitis separata
Stictaspis separata Bezzi, 1913: 104. HT F (ZSI), India. Nagaland: Kohima. Latest references:
Hancock and Drew, 1999: 655; Hancock and McGuire, 2002: 5.
Acroceratitis flava Premlata and Singh, 1988: 635. HT F (PUCP), India. Punjab: Chandigarh.
Synonymy: Hancock and Drew, 1999: 654.
Distribution: India (Punjab, Nagaland), Thailand.
Acroceratitis septemmaculata
Acroceratitis septemmaculata Hardy, 1973: 231. HT F (KUB), Thailand. Phu Kae. Latest reference:
Hancock and Drew, 1999: 655; Hancock and McGuire, 2002: 5.
Distribution: Thailand.
Acroceratitis siamensis
Chelyophora siamensis Munro, 1935a: 17. HT M (ZSI), Thailand. Nakhon Ratchasima: Kao, Lat Bua
[Lat Bua Khao Station, 14º52'N 101º36'E]. Latest reference: Hancock and Drew, 1999: 656.
Acroceratitis similis Hardy, 1973: 233. HT M (KUB), Thailand. Phu Kae. Synonymy:
Hancock and Drew, 1999: 656.
Distribution: Thailand.
Acroceratitis striata
Ceratitis striata Froggatt, 1909: 111. LT M (NSWA), Sri Lanka: Central: Peradeniya, Royal Botanic
Gardens. Lectotype designated by Hancock and Drew (1999: 658).
Distribution: Sri Lanka.
Acroceratitis tenmalaica
Acroceratitis tenmalaica Hancock and Drew, 1999: 659. HT M (BMNH), India. Kerala: Tenmalai,
500-800 ft.
Distribution: India (Kerala).
Acroceratitis tomentosa
Acroceratitis tomentosa Hardy, 1973: 235. HT F (KUB), Thailand. Bangkok, Bangkhen. Latest
reference: Hancock and Drew, 1999: 661.
Acroceratitis maculata Premlata and Singh, 1988: 635. HT M (PUCP?), India. Punjab: Chandigarh.
Synonymy: Hancock and Drew, 1999: 661.
Distribution: India (Punjab), Thailand.
2. ACROTAENIOSTOLA
Acrotaeniostola Hendel, 1914: 80. Type species: sexvittata Hendel (OD). Latest reference: Hancock
and Drew, 1999: 662.
Acrotaeniostola apiventris
Acrotaeniostola apiventris Munro, 1935a: 19. HT M (ZSI), India. W. Bengal: e. Himalayas,
Darjeeling Dist., Pashok, 2000 ft. Latest reference: Hancock and Drew, 1999: 664.
Distribution: India (W. Bengal, Arunachal Pradesh, Meghalaya), n. Thailand.
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BIOTAXONOMY OF TEPHRITOIDEA
Acrotaeniostola flavoscutellata
Acrotaeniostola flavoscutellata Shiraki, 1933: 149. HT F (NTU), Taiwan. Shinchiku or Musha.
Described from 2 females from 2 localities, but which is type locality not specified. Latest reference:
Hancock and Drew, 1999: 665.
Acrotaeniostola antennata Shiraki, 1968: 49. HT M (USNM), Japan. Ryukyu Is.: Okinawa I.
Synonymy: Hardy, 1973: 170.
Distribution: Japan (Ryukyu Is.), Taiwan.
Acrotaeniostola fuscinotum
Acrotaeniostola fuscinotum Hering, 1938c: 16. HT M (NRS), Burma. Kachin: Kambaiti [25°24'N
98°9'E]. Latest reference: Hancock and Drew, 1999: 666.
Distribution: Burma, Vietnam.
Acrotaeniostola longicauda
Acrotaeniostola longicauda Wang, 1998: 28. HT M (IZAS), China. Yunnan: Menglongbanna (21.5°N
100.6°E), 1600 m. Treated as synonym of A. morosa (Hering), by Hancock and Drew (1999: 774).
Reinstated from synonymy in this publication (see Explanations to taxonomic changes, at the end of this
catalog).
Acrotaeniostola morosa: Hancock and Drew, 1999: 666. Misid. (Thailand specimens).
Distribution: China (Yunnan), n. Thailand.
Acrotaeniostola morosa
Taeniostola morosa Hering, 1938c: 15. HT F (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Distribution: n. Burma.
Acrotaeniostola pieli
Acrotaeniostola pieli Zia, 1937: 157. HT M (IZAS), China. Zhejiang: Tien-Mo-Shan [Tianmushan].
Latest references: Wang, 1998: 29; Hancock and Drew, 1999: 668.
Distribution: China (Zhejiang).
Acrotaeniostola quadrivittata
Acrotaeniostola hoenei ssp. quadrivittata Chen, 1948: 94. HT F (IZAS), China. Fujian: Shao-woo
[Shaowu]. Latest references: Wang, 1998: 29; Hancock and Drew, 1999: 668.
Distribution: China (Hubei, Fujian, Yunnan, Hainan).
Acrotaeniostola quinaria
Trypeta quinaria Coquillett, 1910: 308. ST B (USNM), China. Hong Kong. Latest references: Wang,
1998: 29; Hancock and Drew, 1999: 669.
Acrotaeniostola rubra Chen, 1948: 95. HT M (IZAS), Vietnam. Hoa-Binh. Synonymy: Hardy, 1973:
171; Hancock and Drew, 1999: 669.
Spilographa quadrifasciata Enderlein, 1911: 436. HT F (PAN), Indonesia. Sumatra: Soekaranda.
Type data: Hardy, 1988: 83. Synonymy: Hancock and Drew, 1999: 669.
Acrotaeniostola quinaria: Chen, 1948: 72.
Distribution: China (Guangdong, Hong Kong, Hainan), Thailand, Laos, Vietnam, Malaysia (w. and
Sabah), Indonesia (Sumatra).
Acrotaeniostola scutellaris
Trypeta scutellaris Matsumura, 1916: 415. ST M (HUS), Japan. Hokkaido: Sapporo; and Honshu:
Towada. Syntypes not located in HUS by Wang (1998: 30). Latest references: Korneyev, 1997: 34; Wang,
1998: 28, 30; Hancock and Drew, 1999: 670.
Acrotaeniostola hoenei Hering, 1936: 57. HT F (BMNH), China. Zhejiang: W. Tien-Mu-Shan
[Tianmushan]. Synonymy: Hancock and Drew 1999: 670.
Acrotaeniostola hönei Hering, 1936: 57, Incosp. hoenei Hering. Automatic correction under Art.
32.5.2.1.
Acrotaeniostola honei Wang, 1998: 28, Missp. hoenei Hering.
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Kovac et al. / Catalog and classification of Gastrozonini
Distribution: China (Zhejiang, Guangxi), Korea, Russia (Kurile Is.), Japan (Hokkaido, Honshu, Shikoku,
Kyushu).
Acrotaeniostola sexvittata
Acrotaeniostola sexvittata Hendel, 1914: 80. T A (MNM, NMW), Formosa [Taiwan]. Latest
references: Wang, 1998: 30, Hancock and Drew, 1999: 672.
Acrotaeniostola helvenaca Ito, 1984a: 71. HT M (UOPJ), Japan. Ryukyu Is.: Iriomote I., Omotoyama.
Synonymy: Hancock and Drew, 1999: 672.
Acrotaeniostola sexvittata Hendel, 1915: 438. ST B (MNM, NMW), Taiwan. Taihorin; Mt. Hoozan;
and Kankau. Preoccupied by Hendel, 1914. Type data: Hardy, 1968: 108.
Acrotaeniostola helvanaca Norrbom et al., 1999: 72, Missp. helvenaca Ito.
Distribution: Japan (Ryukyu Is.), Taiwan.
Acrotaeniostola spiralis
Acrotaeniostola spiralis Munro, 1935a: 18. HT F (ZSI), Bangladesh. Chittagong Hills, Rangamati.
Latest references: Wang, 1998: 30; Hancock and Drew, 1999: 673.
Distribution: Bangladesh, India (Meghalaya), China (Yunnan, Hainan), Laos, Malaysia (Sabah),
Indonesia (Sumatra).
Acrotaeniostola yunnana
Acrotaeniostola yunnana Wang, 1998: 31. HT F (IZAS), China. Yunnan: Cheli (22.0°N 100.8°E),
700 m. Latest reference: Hancock and Drew, 1999: 774, 775.
Distribution: India (Meghalaya), China (Yunnan).
3. ANOPLOMUS
Anoplomus Bezzi, 1913: 100. Type species: flexuosus Bezzi (= cassandra Osten Sacken) (OD). Latest
reference: Hancock, 1999: 936.
Anoplomus cassandra
Trypeta cassandra Osten Sacken, 1882: 228. HT M (DEI), Philippines. Latest references: Wang,
1998: 14; Hancock, 1999: 936.
Anoplomus flexuosus Bezzi, 1913: 100, N. N. fasciventris Macquart, 1848.
Tephritis fasciventris Macquart, 1848: 225. ST B (Payen), Indonesia. Java. Preoccupied by Macquart,
1843.
Distribution: India, China (Yunnan), Burma, Thailand, Laos, Philippines, Indonesia (Java).
Anoplomus hainanensis
Anoplomus hainanensis Wang, 1998: 14. HT M (IZAS), China. Hainan: Yinggen (19.0°N 109.8°E),
200 m.
Distribution: China (Hainan).
Anoplomus nigrifemoratus
Anoplomus nigrifemoratus Hardy, 1973: 242. HT M (BBM), Laos. Vientiane: Ban Van Eue.
Distribution: Laos.
Anoplomus rufipes
Anoplomus rufipes Hardy, 1973: 243. HT M (BBM), Thailand. Chiang Mai: Chiang Dao.
Distribution: Thailand, Laos.
4. BISTRISPINARIA
Bistrispinaria Speiser, 1913: 145. Type species: Ceratitis fortis Speiser (OD). Proposed as a subgenus
of Ceratitis. Latest reference: Hancock, 1999: 914.
Bistrispinaria atlas
Clinotaenia atlas Munro, 1957: 866. HT M (BMNH), Uganda. Ruwenzori Range, Namwamba
Valley, 6500 ft. Latest reference: Hancock, 1999: 915.
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BIOTAXONOMY OF TEPHRITOIDEA
Clinotaenia atlas Munro, 1956: 465. HT F (MRAC?), Burundi. Urundi, Bururi, 1800-2000 m. Nomen
nudum. Published after 1930 without a description or bibliographic reference to one.
Distribution: Uganda, Burundi, Kenya.
Bistrispinaria fortis
Ceratitis fortis Speiser, 1913: 145. HT M (Unknown), Cameroon. Mt. Cameroon, Soppo. Latest
reference: Hancock, 1999: 916.
Pardalaspis aglaspis Séguy, 1941: 117. T M (MNHNP), Ivory Coast. Kouibly. Synonymy: Munro,
1957: 867.
Distribution: Ivory Coast, Nigeria, Cameroon, Zaire, Uganda.
Bistrispinaria magniceps
Chelyophora magniceps Bezzi, 1918: 229. HT F (BMNH), Sudan. Latest reference: Hancock, 1999:
918.
Chelyophora lemniscata Enderlein, 1920: 355. ST B (ZMHU), Kenya. Mombassa; and Tanzania.
Mtoashimu; and Kwasengiwa. Synonymy: Bezzi, 1924b: 98.
Distribution: Sudan, Uganda, Kenya, Tanzania, Malawi, Mozambique.
Bistrispinaria woodi
Chelyophora woodi Bezzi, 1924b: 98. LT M (BMNH), Malawi. Cholo. Lectotype designated by
Hancock (1999: 919).
Chelyophora uranos Hering, 1942: 280. ST B (ZMHU), Cameroon. Uam region, Bosum. Synonymy:
Hancock, 1999: 919.
Chelyophora woodi Bezzi, 1924a: 14. HT M (MRAC), “Congo da Lemba”. Nomen nudum. Published
without a diagnosis or indication; the only character mentioned is said to be shared by another species
(Hancock, 1999: 921).
Distribution: Nigeria, Cameroon, Zaire, Rwanda or Burundi (“Ruanda-Urundi”, as per Munro 1956: 465),
Malawi.
5. CARPOPHTHORELLA
Carpophthorella Hendel, 1914: 80. Type species: magnifica Hendel (OD). Latest reference: Hancock
and Drew, 1999: 673.
Carpophtherella Hardy, 1974: 155, Missp. Carpophthorella Hendel.
Carpophthorella capillata
Gastrozona capillata Bezzi, 1914: 324. ST F (Baker), Philippines. Luzon, Laguna: Los Banos.
Lectotype designation of Hardy (1969: 481) invalid; syntypes currently in MCSNM. Latest reference:
Hancock and Drew, 1999: 675.
Gastrozona luteiseta Bezzi, 1914: 325. ST M (Baker), Philippines. Luzon, Laguna: Los Banos.
Lectotype designation of Hardy (1969: 481) invalid; syntypes currently in MCSNM. Synonymy: Hancock
and Drew, 1999: 676.
Distribution: Philippines, ?Malaysia (Sabah) [possible misidentification of Carpophthorella sookae].
Carpophthorella magnifica
Carpophthorella magnifica Hendel, 1914: 80. T A (MNM), Formosa [Taiwan]. Latest reference:
Hancock and Drew, 1999: 677.
Carpophthorella magnifica Hendel, 1915: 449. ST B (MNM), Taiwan. Kankau. Preoccupied by
Hendel, 1914.
Distribution: Taiwan.
Carpophthorella nigrifascia
Trypeta nigrifascia Walker, 1860: 158. LT M (BMNH), Indonesia. Sulawesi: Makassar [Ujung
Padang]. Lectotype designated by inference of holotype by Hardy (1959: 218). Latest reference: Hancock
and Drew, 1999: 678.
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Gastrozona albiscutellata Enderlein, 1920: 354. HT F (ZMHU), Indonesia. Sumatra: Padang,
Bungus-Bucht [Bungus-Brucht]. Type data and Synonymy: Hardy, 1988: 90.
Carpophthorella setifrons Malloch, 1939: 263. HT F (BMNH), Solomon Is. Guadalcanal. Synonymy:
Permkam and Hancock, 1995: 1328.
Trypeta retorta Walker, 1861: 16. LT F (BMNH), Indonesia. Maluku: Gilolo [Djailolo]. Lectotype
designated by inference of holotype by Hardy (1959: 220). Synonymy: Hardy, 1988: 90.
Gastrozona bifasciata Meijere, 1916: 48. HT F (ZMAN), Indonesia. Sumatra: Simalur I., Labuan
Badjau. Type data and Synonymy: Hardy, 1988: 90.
Distribution: Indonesia (Sumatra, Sulawesi, Maluku), Malaysia (Sarawak), Papua New Guinea, Solomon
Is., Australia (Queensland).
Carpophthorella semipennata
Carpophthorella semipennata Hering, 1938c: 9. HT M (NRS), Burma. Kachin: Kambaiti [25°24'N
98°9'E]. Latest reference: Hancock and Drew, 1999: 679.
Distribution: Burma.
Carpophthorella sookae
Carpophthorella sookae Chua, 2003: 465. HT M (BMNH), Malaysia: Ulu Langat, Selangor.
Distribution: w. Malaysia.
Carpophthorella semipennata: Hancock and Drew, 1999: 679 [misidentification (Malaysian female),
see Chua (2003: 468)].
6. CERATITOIDES
Ceratitoides Hendel, 1928: 365. Type species: nigromaculata Hendel (OD). Latest reference:
Hancock, 1999: 939.
Ceratitoides nigromaculata
Ceratitoides nigromaculata Hendel, 1928: 366. HT F (DEI), Uganda. Latest reference: Hancock,
1999: 940.
Chelyophora frigida Hering, 1942: 281. HT M (ZMHU), Cameroon. Jaunde [Yaounde] region.
Synonymy: Hancock, 1999: 940.
Distribution: Cameroon, Uganda.
7. CHAETELLIPSIS
Chaetellipsis Bezzi, 1913: 126. Type species: paradoxa Bezzi (OD). Latest reference: Hancock and
Drew 1999: 681.
Podophysa Hering, 1938c: 8. Type species: pretiosa Hering (OD). Synonymy: Hancock, 1991: 122.
Poecillis Bezzi, 1913: 128. Type species: judicanda Bezzi (= paradoxa Bezzi) (OD). Synonymy:
Hardy, 1973: 177.
Chaetellipsis alternata
Podophysa alternata Zia, 1963: 457. HT M (IZAS), China. Yunnan: Xi-Sang-Ban-Na
[Xishuangbanna]. Latest reference: Hancock and Drew, 1999: 683.
Distribution: China (Yunnan), n. Thailand.
Chaetellipsis bivittata
Carpophthorella bivittata Hardy, 1988: 89. HT F (BBM), Malaysia. Sabah: 19 km N. of Kalabakan,
forest camp, 60 m. Latest references: Chua, 1999: 196; Hancock and Drew 1999: 684.
Distribution: s. Thailand, Malaysia (w., Sarawak, Sabah).
Chaetellipsis dispilota
Chaetellipsis dispilota Hardy, 1973: 180. HT M (BBM), India. Uttar Pradesh: Ranikhet. Latest
reference: Hancock and Drew, 1999: 687. Considered synonym of C. alternata by Wang (1998: 33), but
possibly only paratypes from Thailand are that species (Hancock and Drew, 1999: 683).
Distribution: India (Uttar Pradesh).
179
BIOTAXONOMY OF TEPHRITOIDEA
Chaetellipsis kinabaluensis
Chaetellipsis kinabaluensis Hancock and Drew, 1999: 689. HT M (BMNH), Malaysia. Sabah: Poring
Hot Springs, nr. Mt. Kinabalu.
Distribution: Malaysia (Sabah).
Chaetellipsis maculosa
Chaetellipsis maculosa Hancock and Drew, 1999: 690. HT M (BMNH), Malaysia. Selangor: Ulu
Langat.
Distribution: w. Malaysia (Selangor), Indonesia (Java).
Chaetellipsis paradoxa
Chaetellipsis paradoxa Bezzi, 1913: 127. HT M (ZSI), India. Bihar: Parasnath, 4400 ft. Latest
reference: Hancock and Drew, 1999: 693.
Distribution: Pakistan, India, Sri Lanka, China (Yunnan), Burma, Thailand, Laos.
Poecillis judicanda Bezzi, 1913: 128. ST F (ZSI), India. Bihar: Parasnath, 4300 ft. Synonymy: Hardy,
1973: 182.
Chaetellipsis atrata Hardy, 1973: 179. HT F (BBM), Laos. Vientiane: Ban Van Eue. Synonymy:
Hancock and Drew, 1999: 693.
Gastrozona flavostriata Hering, 1938c: 12. HT F (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Synonymy: Hardy, 1973: 182.
Podophysa occipitalis Zia, 1963: 459. HT M (IZAS), China. Yunnan: Xi-Sang-Ban-Na
[Xishuangbanna]. Synonymy: Hancock and Drew, 1999: 693.
Chaetellipsis pretiosa
Podophysa pretiosa Hering, 1938c: 9. HT M (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Latest reference: Hancock and Drew, 1999: 695.
Distribution: Burma.
8. CHELYOPHORA
Chelyophora Rondani, 1875: 433. Type species: borneana Rondani (MO). Latest reference: Hancock
and Drew, 1999: 697.
Chelyophora borneana
Chelyophora borneana Rondani, 1875: 434. LT F (MCSNG), Malaysia. Sarawak. Lectotype
designated by inference of holotype by Hardy (1988: 92). Latest reference: Hancock and Drew, 1999: 697.
Distribution: Malaysia (Sarawak).
9. CLINOTAENIA
Clinotaenia Bezzi, 1920: 225. Type species: anastrephina Bezzi (OD). Latest reference: Hancock,
1999: 921.
Clinotaenia anastrephina
Clinotaenia anastrephina Bezzi, 1920: 226. HT F (BMNH), Malawi. Mlanje: Mt. Mlanje [Sapitwa].
Latest reference: Hancock, 1999: 922.
Distribution: Zaire, Malawi.
Clinotaenia camerunica
Clinotaenia camerunica Hancock, 1999: 923. HT F (BMNH), Cameroon. Uam region, near Bosum.
Distribution: Cameroon.
Clinotaenia grata
Trypeta grata Wiedemann, 1830: 498. T F (ZMHU), Kap [South Africa. Cape Province or Cape of
Good Hope]. Latest reference: Hancock, 1999: 925.
Clinotaenia cedarensis Munro, 1933: 30. HT F (SANC), South Africa. Natal: Cedara. Synonymy:
Hancock, 1999: 925.
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Distribution: South Africa (e. Cape, Natal).
Clinotaenia inyanga
Clinotaenia inyanga Hancock, 1985: 62. HT F (NMBZ), Zimbabwe. Nyanga, Rhodes Inyanga
Orchard. Latest reference: Hancock, 1999: 927.
Distribution: Zaire, Zimbabwe.
Clinotaenia superba
Carpophthoromyia superba Bezzi, 1918: 226. HT F (BMNH), Malawi. Limbe, Chiromo, Ruo R.
Latest reference: Hancock 1999: 928.
Distribution: Malawi, Mozambique.
10. CYRTOSTOLA
Cyrtostola Hancock and Drew, 1999: 699. Type species: Taeniostola limbata Hendel (OD).
Cyrtostola limbata
Taeniostola limbata Hendel, 1915: 435. ST B (DEI, MNM, BMNH), Taiwan. Taitorinsho; Taihorin;
Sokutsu. Also possible syntypes in NMW (Hardy, 1968: 124). Latest reference: Hancock and Drew,
1999: 699.
Distribution: India (Uttar Pradesh, Assam), Nepal, China (Yunnan), Burma, Thailand, w. Malaysia,
Taiwan; Papua New Guinea? [error?].
11. DIETHERIA
Dietheria Hardy, 1973: 183. Type species: fasciata Hardy (OD). Latest reference: Hancock and Drew,
1999: 701.
Dietheria fasciata
Dietheria fasciata Hardy, 1973: 184. HT M (BBM), Vietnam. Ban Me Thuot, 500 m. Latest reference:
Hancock and Drew, 1999: 701.
Distribution: Thailand, China (Yunnan), Vietnam.
Dietheria gracilis
Taeniostola gracilis Bezzi, 1913: 120. HT F (ZSI), Burma. Karen: base of Dawna Hills. Latest
reference: Hancock and Drew, 1999: 702.
Distribution: Burma.
12. ENICOPTERA
Enicoptera Macquart, 1848: 223. Type species: flava Macquart (OD). Latest reference: Hancock and
Drew, 1999: 702.
Henicoptera Loew, 1873: 21, Emend. Enicoptera Macquart.
Enicoptera cuneilineata
Henicoptera cuneilineata Hering, 1937: 108. HT M (BMNH), Philippines. Luzon, Manila. Latest
reference: Hancock and Drew, 1999: 706.
Enicoptera cuneilinea Hardy, 1974: 166, Missp. cuneilineata Hering.
Distribution: Philippines (Luzon).
Enicoptera flava
Enicoptera flava Macquart, 1848: 223. HT M (Payen), Indonesia. Java. Holotype stated to be in
MNHNP, but not examined (Hancock and Drew, 1999: 706).
Distribution: Indonesia (Java).
Enicoptera flavofemoralis
Henicoptera flavofemoralis Hering, 1937: 107. HT M (BMNH), Philippines. Luzon, Bataan: Limay
[14°34'N 120°36'E]. Latest reference: Hancock and Drew, 1999: 707.
Distribution: Philippines (Luzon).
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BIOTAXONOMY OF TEPHRITOIDEA
Enicoptera gigantea
Enicoptera gigantea Enderlein, 1911: 413. LT M (PAN), Indonesia. Sumatra: Soekaranda. Lectotype
designated by Hardy (1969: 480). Latest reference: Hancock and Drew, 1999: 708.
Distribution: w. Malaysia, Indonesia (Sumatra).
Enicoptera gressitti
Enicoptera gressitti Hardy, 1988: 97. HT F (BBM), Malaysia. Sabah: Sandakan Bay (SW.), Sapagaya
lumber camp, 2-20 m. Latest reference: Hancock and Drew, 1999: 709.
Distribution: Malaysia (Sabah, Sarawak).
Enicoptera il
Henicoptera il Hering, 1938b: 411. HT M (BMNH), Philippines. Luzon. Latest reference: Hancock
and Drew, 1999: 711.
Distribution: Philippines (Luzon, Leyte).
Enicoptera interrupta
Henicoptera interrupta Hering, 1937: 107. HT M (BMNH), Philippines. Mindanao, Surigao del
Norte: Surigao. Latest reference: Hancock and Drew, 1999: 712.
Distribution: Philippines (Mindanao).
Enicoptera palawanica
Henicoptera palawanica Hering, 1942: 278. HT F (ZMHU), Philippines. Palawan. Latest reference:
Hancock and Drew 1999: 713.
Distribution: Philippines (Palawan).
Enicoptera proditrix
Enicoptera proditrix Osten Sacken, 1882: 233. LT M (DEI), Philippines. Lectotype designated by
Hardy (1969: 478). Latest reference: Hancock and Drew 1999: 714.
Distribution: Philippines.
Enicoptera spoliata
Henicoptera spoliata Hering, 1937: 106. HT M (BMNH), Philippines. Mindanao, Zamboanga del
Sur: Port Banga [7°30'N 122°26'E]. Latest reference: Hancock and Drew, 1999: 715.
Distribution: Philippines (Mindanao).
Enicoptera sumatrana
Henicoptera sumatrana Hering, 1938b: 412. ST B (PAN, BMNH), Indonesia. Sumatra: Soekaranda.
Latest reference: Hancock and Drew, 1999: 716.
Enicoptera proditrix Enderlein 1911: 414, Misid.
Distribution: Indonesia (Sumatra, Java, Sumbawa).
Enicoptera tortuosa
Enicoptera tortuosa Walker, 1860: 155. LT M (BMNH), Indonesia. Sulawesi: Makassar
[Ujung Padang]. Lectotype designated by inference of holotype by Hardy (1959: 188). Latest reference:
Hancock and Drew, 1999: 717.
Distribution: Indonesia (Sulawesi).
13. GALBIFASCIA
Galbifascia Hardy, 1973: 247. Type species: sexpunctata Hardy (OD). Latest reference: Hancock and
Drew 1999: 718.
Galbifascia sexpunctata
Galbifascia sexpunctata Hardy, 1973: 248. HT M (BBM), Laos. Vientiane: Muong Tourakom, 120 m.
Latest reference: Hancock and Drew, 1999: 719.
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Galbifascia quadripunctata Hardy, 1973: 247. HT F (BMNH), Sri Lanka. Central: Pundaluoya
[7°01'N 80°40'E]. Synonymy: Hancock and Drew 1999: 719.
Distribution: India (Kerala), Sri Lanka, China (Yunnan), Thailand, Laos, Vietnam, Philippines (Luzon).
14. GASTROZONA
Gastrozona Bezzi, 1913: 105. Type species: Tephritis fasciventris Macquart (OD). Latest reference:
Hancock and Drew, 1999: 71.
Gastrozoa Shinji, 1939: 289, Missp. Gastrozona Bezzi.
Gastrozona balioptera
Gastrozona balioptera Hardy, 1973: 188. HT M (BMNH), Thailand. Chiang Mai: Chiang Dao. Latest
reference: Hancock and Drew, 1999: 722.
Distribution: India (Arunachal Pradesh), China (Yunnan), Burma, Thailand.
Gastrozona fasciventris
Tephritis fasciventris Macquart, 1843: 382. T M (MNHNP), “Indes orientales”. Latest reference:
Hancock and Drew, 1999: 723.
Gastrozona melanista Bezzi, 1913: 107. HT F (ZSI), India. Kerala: on ship, 5 mi. off Calicut
[Kozhikode], Malabar Coast. Synonymy: Hardy, 1973: 190.
Gastrozona macquarti Hendel, 1913a: 38. ST B (DEI, NMW), Taiwan. Kanshirei. Type data: Hardy,
1968: 117. Synonymy: Hardy, 1973: 190.
Gastrozona melanophila Hering, 1940a: 3. HT M (BMNH), Taiwan. Tao Tsui Kutsu. Synonymy:
Hardy, 1973: 190.
Tephritis vittata Macquart, 1851: 263. T F (MNHNP), Asia. Synonymy: Hardy, 1973: 190.
Gastrozona appendiculata Zia, 1938: 22. HT M (IZAS), China. se. Gansu: Cheumen [Yumen].
Synonymy: Hardy, 1973: 190.
Tephritis fusciventris Macquart, 1843: 459, Incosp. fasciventris Macquart. Hardy, 1973: 190 (FR).
Distribution: China (Gansu), India, Bangladesh, Burma, Thailand, Laos, Vietnam, w. Malaysia, Indonesia
(Sumatra), Taiwan.
Gastrozona hirtiventris
Gastrozona hirtiventris Chen, 1948: 97. HT M (IZAS), China. Zhejiang: Mokanshan [Moganshan].
Latest references: Wang, 1998: 37, Hancock and Drew 1999: 722, Hancock and McGuire, 2002: 6.
Distribution: China (Zhejiang), Thailand.
Gastrozona isis
Gastrozona isis Hering, 1938c: 12. HT M (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E]. Latest
reference: Hancock and Drew, 1999: 726.
Distribution: Burma.
Gastrozona montana
Gastrozona montana Bezzi, 1913: 106. ST B (ZSI), India. W. Bengal: e. Himalayas, Kurseong, 5000
ft. Latest reference: Hancock and Drew, 1999: 728.
Distribution: India (Assam, W. Bengal), Burma.
Gastrozona parviseta
Gastrozona parviseta Hardy, 1973: 192. HT M (BBM), Thailand. Chiang Mai: Chiang Dao, 450 m.
Latest reference: Hancock and Drew, 1999: 729.
Gastrozona menglanica Wang, 1998: 37. HT F (BAUC), China. Yunnan: Mengla (21.4°N 101.5°E),
800 m. Synonymy: Hancock and Drew, 1999: 774.
Distribution: India (Karnataka), China (Yunnan), Burma, Thailand.
Gastrozona proterva
Gastrozona proterva Hering, 1938c: 13. HT M (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Latest references: Hardy, 1973: 187; Hancock and Drew, 1999: 730.
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BIOTAXONOMY OF TEPHRITOIDEA
Gastrozona solitaria Hering, 1938c: 14. HT F (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Synonymy: Hancock and Drew, 1999: 730.
Distribution: India, Burma.
Gastrozona quadrivittata
Gastrozona quadrivittata Wang, 1992: 1150. HT F (IZAS), China. Guizhou: Mt. Leigong (26.4°N
108.2°E), 1700-2000 m.
Distribution: China (Guizhou, Hunan).
Gastrozona selangorensis
Gastrozona selangorensis Chua, 2003: 468. HT M (BMNH), Malaysia (Ulu Langat).
Distribution: Malaysia (Peninsular).
Gastrozona soror
Acidia soror Schiner, 1868: 264. HT M (NMW), Indonesia. Java: Batavia [Jakarta]. Type data: Hardy,
1968: 141.
Gastrozona fasciventris Bezzi, 1913: 105, Misid. See Hancock and Drew (1999: 732).
Distribution: India (Assam), Thailand, Indonesia (Java).
15. ICHNEUMONOPSIS
Ichneumonopsis Hardy, 1973: 132. Type species: burmensis Hardy (OD).
Ichneumonopsis burmensis
Ichneumonopsis burmensis Hardy, 1973: 133. HT M (BMNH), Burma. Chin: Chin Hills, Mount
Victoria [21°14'N 93°55'E], 1400 m. Latest reference: Drew and Hancock, 1994: 830; Radhakrishnan,
2000: 203.
Distribution: India (Meghalaya), n. Burma.
16. LEUCOTAENIELLA
Leucotaeniella Bezzi, 1918: 227. Type species: trispila Bezzi (OD). Latest reference: Hancock, 1999:
930.
Leucotaeniella guttipennis
Leucotaeniella guttipennis Bezzi, 1920: 223. HT F (BMNH), Nigeria. Zungeru. Latest references:
Hancock, 1999: 930; Hancock et al., 2001: 49.
Distribution: Guinea, Nigeria, Zaire, Uganda, Zambia, Angola.
Leucotaeniella mambillae
Leucotaeniella mambillae Hancock, 1999: 932. HT F (BMNH), Nigeria. Mambilla Plateau, Ngel
Nyaki.
Distribution: Nigeria.
Leucotaeniella pentaspila
Leucotaeniella pentaspila Bezzi, 1918: 229. HT F (BMNH), Sudan. Latest reference: Hancock, 1999:
933.
Distribution: Sudan, Zaire, Angola, Zambia.
Leucotaeniella trispila
Leucotaeniella trispila Bezzi, 1918: 228. LT M (BMNH), Malawi. Limbe, Chiromo, Ruo R.
Lectotype designated by Hancock (1999: 935).
Distribution: Zambia, Malawi, Zimbabwe.
17. PARAGASTROZONA
Paragastrozona Shiraki, 1933: 154. Type species: Gastrozona japonica Miyake (OD). Latest
reference: Hancock and Drew, 1999: 733.
Paragastrozoa Shinji, 1940a: 162, Missp. Paragastrozona Shiraki.
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Paragastrozona apicemaculata
Gastrozona apicemaculata Hering, 1938c: 11. HT F (NRS), Burma. Kachin: Kambaiti [25°4'N
98°9'E]. Latest reference: Hancock and Drew, 1999: 734.
Distribution: Burma.
Paragastrozona fukienica
Gastrozona fukienica Hering, 1953: 5. HT F (ZFMK), China. Fujian: Kuatun. Latest reference:
Hancock and Drew, 1999: 737.
Distribution: China (Fujian).
Paragastrozona japonica
Gastrozona japonica Miyake, 1919: 152. ST B (Unknown), Japan. Honshu: near Tokyo, Oji.
Syntypes apparently lost (Shiraki, 1933: 156).
Gastrozona japonica var. miyakei Bezzi, 1926: 265. ST A (MCSNM), Japan. Hokkaido: Sapporo.
Distribution: e. Russia, Korea, Japan (Hokkaido, Honshu, Shikoku, Kyushu), Taiwan.
Paragastrozona orbata
Gastrozona orbata Hering, 1938c: 10. HT F (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Latest reference: Hancock and Drew, 1999: 740.
Distribution: Burma.
Paragastrozona quinquemaculata
Paragastrozona quinquemaculata Wang, 1998: 40. HT M (IZAS), China. Sichuan: Mt. Emei (29.5°N
103.3°E). Latest reference: Hancock and Drew, 1999: 736.
Distribution: China (Sichuan).
Paragastrozona tripunctata
Taeniostola tripunctata Shiraki, 1968: 52. HT M (USNM), Japan. Ryukyu Is. Latest reference:
Hancock and Drew, 1999: 741.
Gastrozona tripunctata: Hardy, 1973: 186.
Acroceratitis tripunctata: Hardy, 1977: 88.
Paragastrozona tripunctata: Ito, 1984b: 135.
Distribution: Japan (Ryukyu Is.).
Paragastrozona trivittata
Paragastrozona trivittata Hancock and Drew, 1999: 742. HT F (BMNH), Burma. Adung Valley,
8000 ft.
Distribution: n. Burma.
Paragastrozona vulgaris
Gastrozona vulgaris Zia, 1937: 151. HT M (IZAS), China. Jiangsu: Nanking [Nanjing]. Sex of
holotype and type locality not indicated in original description, but determined from label data by Wang
(1998: 39). Latest reference: Hancock and Drew, 1999: 744.
Taeniostola melli Hering, 1942: 276. HT M (ZMHU), China. Guangdong: Canton [Guangzhou].
Synonymy: Wang, 1998: 39; Hancock and Drew, 1999: 744.
Distribution: China (Sichuan, Anhui, Jiangsu, Zhejiang, Hunan, Fujian, Guangdong).
18. PARAXARNUTA
Paraxarnuta Hardy, 1973: 195. Type species: bambusae Hardy (OD). Latest reference: Hancock and
Drew, 1999: 745.
Paraxarnuta anephelobasis
Paraxarnuta anephelobasis Hardy, 1973: 196. HT M (BBM), Thailand. Loei: 12-15 km NW. of Loei,
275 m. Latest references: Wang, 1998: 42; Hancock and Drew, 1999: 746.
Distribution: China (Yunnan), Thailand.
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BIOTAXONOMY OF TEPHRITOIDEA
Paraxarnuta bambusae
Paraxarnuta bambusae Hardy, 1973: 197. HT M (BBM), Laos. Vientiane: Muong Tourakom, 180 m.
Latest references: Wang 1998: 42; Hancock and Drew, 1999: 746.
Distribution: e. India, China (Yunnan), Thailand, Laos, Vietnam.
Paraxarnuta extorris
Acrotaeniostola extorris Hering, 1942: 277. HT F (ZMHU), Type locality unknown, probably
“Indien” [Indonesia?]. Latest reference: Hancock and Drew, 1999: 748.
Distribution: w. Malaysia (Selangor), Indonesia (Java).
Paraxarnuta interrupta
Acrotaeniostola interrupta Hardy, 1988: 82. HT M (BBM), Malaysia. Selangor: Ulu Langat, 300390 m. Latest reference: Hancock and Drew, 1999: 750.
Distribution: w. Malaysia.
Paraxarnuta maculata
Paraxarnuta maculata Wang, 1998: 42. HT M (IZAS), China. Yunnan: Cheli (22.0°N 100.8°E), 700 m.
Distribution: China (Yunnan).
19. PARDALASPINUS
Pardalaspinus Hering, 1952: 282. Type species: Pardalaspis migrata Hering (= laqueata Enderlein)
(OD).
Notophosa Zia, 1964: 53. Type species: connexa Zia (= laqueata Enderlein) (OD). Synonymy:
Hancock and Drew, 1994: 875.
Ceratitisoma Zia, 1964: 54. Type species: bimaculatum Zia (OD). Synonymy: Hancock and Drew,
1994: 875.
Notophysa Hancock and Drew, 1994: 875, Missp. Notophosa Zia.
Pardalaspinus aberratus (N. Comb.)
Acroceratitis aberrata Hardy, 1973: 220. HT M (BBM), Laos. Vientiane: Ban Van Eue. Latest
reference: Hancock, 1999: 946.
Ceratitoides aberratus: Hancock, 1999: 946.
Distribution: Laos.
Pardalaspinus adnatus (N. Comb.)
Acroceratitis adnata Hardy, 1973: 220. HT M (BBM), Laos. Vientiane: Ban Van Eue. Latest
reference: Hancock, 1999: 946.
Ceratitoides adnatus: Hancock, 1999: 946.
Distribution: Laos.
Pardalaspinus adversarius
Pardalaspinus adversarius Hering, 1952: 283. HT M (RNH), Indonesia. w. Java: Radjamandala, 2300 m. Latest reference: Hancock, 1999: 940.
Ceratitoides adversarius: Hancock, 1999: 940.
Distribution: Burma, Indonesia (Java).
Pardalaspinus bimaculatus
Ceratitisoma bimaculatum Zia, 1964: 50. HT M (IZAS), China. Yunnan: Shishong-Baanna
[Xishuangbanna], Damenglung, 650 m. Latest reference: Hancock, 1999: 940.
Proanoplomus minor Hardy, 1973: 270. HT M (BBM), Thailand. Chiang Mai: Chiang Dao, 450 m.
Pardalaspinus bimaculatus: Hancock and Drew, 1994: 876.
Ceratitoides bimaculatus: Hancock, 1999: 940.
Distribution: China (Yunnan, Hainan), Thailand.
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Pardalaspinus laqueatus
Ceratitis laqueata Enderlein, 1920: 347. HT F (ZMHU), Indonesia. Java. Type data:Hardy, 1988: 109.
Latest reference: Hancock, 1999: 946.
Pardalaspis migrata Hering, 1944: 5. HT M (NMW), Ost-Indie [probably Indonesia]. Synonymy:
Hancock and Drew, 1994: 876; Hancock, 1999: 946.
Notophosa connexa Zia, 1964: 49. HT M (IZAS), China. Yunnan: Shishong-Baanna [Xishuangbanna],
Siao-meng-yan, 850 m. Synonymy: Hancock and Drew, 1994: 876; Hancock, 1999: 946.
Pardalaspinus laqueatus: Hancock and Drew, 1994: 876.
Ceratitoides laqueatus: Hancock, 1999: 940.
Distribution: e. India, China (Yunnan), Laos, Vietnam, Indonesia (Java).
Pardalaspinus maai (N. Comb.)
Chelyophora maai Chen, 1948: 92. HT M (IZAS), China. Fujian: Shao-Woo [Shaowu]. Latest
references: Wang, 1998: 25; Hancock, 1999: 946.
Acroceratitis maai: Wang, 1998: 25.
Ceratitoides maai: Hancock, 1999: 939.
Distribution: India (Sikkim), China (Fujian), Laos.
Pardalaspinus namtamai (N. Comb.)
Ceratitoides namtamai Hancock, 1999: 942. HT M (BMNH), Burma. Nam Tamai Valley (27°42'N
97°54'E), 5000 ft.
Distribution: Burma.
Pardalaspinus nitidus
Proanoplomus nitidus Hardy, 1973: 271. HT M (KUB), Thailand. Uthai Thani: Uthai Thani. Latest
reference: Hancock, 1999: 940.
Pardalaspinus nitidus: Hancock and Drew, 1994: 877.
Ceratitoides nitidus: Hancock, 1999: 940.
Distribution: Thailand.
Pardalaspinus sikhimensis (N. Comb.)
Ceratitoides sikhimensis Hancock, 1999: 944. HT M (BMNH), India. Sikkim: Singhik, 5000 ft.
Distribution: India (Sikkim).
Pardalaspinus vittatus
Proanoplomus vittatus Hardy, 1973: 276. HT F (KUB), Thailand. Kanchanaburi: Kanchanaburi.
Latest reference: Hancock, 1999: 940.
Pardalaspinus vittatus: Hancock and Drew, 1994: 878.
Ceratitoides vittatus: Hancock, 1999: 940.
Distribution: India (Sikkim), Burma, Thailand.
Pardalaspinus yongi
Pardalaspinus yongi Hancock and Drew, 1995: 61. HT M (BMNH), Malaysia. Selangor: Kg. Perdik,
Ulu Langat.
Ceratitoides yongi: Hancock, 1999: 940.
Distribution: w. Malaysia.
20. PHAEOSPILA
Phaeospila Bezzi, 1913: 117. Type species: varipes Bezzi (OD). Latest reference: Hancock and Drew,
1999: 751.
Phaeospila dissimilis
Acrotaeniostola dissimilis Zia, 1937: 159. HT M (IZAS), China. Szechuan [Sichuan]: Beibei. Sex of
holotype and type locality not indicated in original description, but determined from label data by Wang
(1998: 27). Latest references: Wang, 1998: 27; Hancock and Drew, 1999: 752.
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BIOTAXONOMY OF TEPHRITOIDEA
Distribution: China (Hubei, Sichuan, Yunnan), Vietnam.
Phaeospila megispilota
Acrotaeniostola megispilota Hardy, 1974: 155. HT F (BBM), Philippines. Luzon, Mountain province:
Abatan, Buguias, 60 km S. of Bontoc, 1800-2000 m. Latest reference: Hancock and Drew, 1999: 754.
Phaeospila megaspilota Hancock and Drew, 1999: 754, Missp. megispilota Hardy.
Distribution: Philippines (Luzon).
Phaeospila varipes
Phaeospila varipes Bezzi, 1913: 118. ST F (ZSI), India. W. Bengal: Darjeeling, 6000 and 7000 ft.
Latest reference: Hancock and Drew, 1999: 754.
Distribution: India (W. Bengal).
21. PHAEOSPILODES
Phaeospilodes Hering, 1939: 170. Type species: torquata Hering (= fenestella Coquillett) (OD).
Latest reference: Hancock and Drew, 1999: 754.
Phaeospilodes bambusae
Phaeospilodes bambusae Hering, 1940b: 322. LT F (BMNH), India. Tamil Nadu: Coimbatore.
Lectotype designated by Hancock and Drew (1999: 755).
Distribution: India (Tamil Nadu).
Phaeospilodes fenestella
Oxyphora fenestella Coquillett, 1910: 308. HT F (USNM), China. Hong Kong. Latest references:
Wang, 1998: 44; Hancock and Drew, 1999: 756.
Phaeospilodes torquata Hering, 1939: 171. HT F (MNHNP), Vietnam. Choganh. Synonymy:
Hancock and Drew, 1999: 756.
Phaeospilodes atrifacies Hering, 1941a: 32. HT M (DEI), Indonesia. Nusa Tenggara: Lombok,
Selong. Synonymy: Hancock and Drew, 1999: 756.
Ptilona poeciloptera Kertész, 1912: 543. ST M (MNM), China. Guangdong: Swatow [Shantou].
Synonymy: Hancock and Drew, 1999: 756.
Oxyna fenestrella Hardy, 1977: 126, Missp. fenestella Coquillett.
Phaeospilodes poeciloptera: Wang, 1998: 44.
Distribution: China (Fujian, Guangxi, Guangdong, Hong Kong, Hainan), Thailand, Vietnam, Indonesia
(Java, Lombok).
Phaeospilodes paragoga
Taeniostola paragoga Hering, 1938c: 16. HT F (NRS), Burma. Kachin: Kambaiti [25°24'N 98°9'E].
Latest reference: Hancock and Drew, 1999: 758, 775.
Taeniostola paragoda Hardy, 1973: 210, Missp. paragoga Hering.
Distribution: e. India (Meghalaya), Burma.
22. PROANOPLOMUS
Proanoplomus Shiraki, 1933: 127. Type species: japonicus Shiraki (OD). Latest reference: Hancock
1999: 937.
Paranoplomus Shiraki, 1933: 131. Type species: formosanus Shiraki (OD).
Parnaoplomus Hardy, 1977: 96, Missp. Paranoplomus Shiraki. Attributed to “authors”.
Proanoplomus affinis
Proanoplomus affinis Chen, 1948: 89. HT F (IZAS), China. Zhejiang: Tianmushan.
Distribution: China (Zhejiang).
Proanoplomus arcus
Paranoplomus arcus Ito, 1949: 53. HT F (UOPJ), Japan. Honshu: Wakayama Prov., Koyasan, 900 m.
Distribution: Japan (Honshu).
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Proanoplomus caudatus
Anoplomus caudatus Zia, 1964: 44. HT F (IZAS), China. Yunnan: Shishong-Baanna
[Xishuangbanna], 700 m.
Distribution: China (Yunnan).
Proanoplomus cinereofasciatus (N. Comb.)
Carpophthoromyia cinereofasciata Meijere, 1924: 37. ST B (ZMAN), Indonesia. Sumatra: Tand
Andalas. Type data: Hardy, 1988: 108. Latest reference: Hancock, 1999: 940.
Ceratitoides cinereofasciatus: Hancock, 1999: 940.
Distribution: Indonesia (Sumatra, Java), Malaysia (Sabah).
Proanoplomus cylindricus
Proanoplomus cylindricus Chen, 1948: 91. HT F (IZAS), Formosa [Taiwan].
Distribution: Taiwan.
Proanoplomus formosanus
Paranoplomus formosanus Shiraki, 1933: 131. ST B (NTU), Taiwan. Arisan. Latest reference:
Hancock, 1999: 938.
Distribution: Taiwan.
Proanoplomus intermedius
Proanoplomus intermedius Chen, 1948: 91. HT F (IZAS), China. Fujian: Shao-Woo [Shaowu].
Distribution: China (Fujian).
Proanoplomus japonicus
Proanoplomus japonicus Shiraki, 1933: 128. ST B (NTU), Japan. Nagano; Kogota; Otoineppu; and
Fukuoka. Latest reference: Hancock, 1999: 937.
Paragastrozoa nigricaudus Shinji, 1940a: 162. HT F (Shinji), Japan. Honshu: near Morioka,
Kamiyonai.
Paragastrozoa nigricaudus Shinji, 1940b: 194. HT F (Shinji), Japan. Honshu: near Morioka.
Preoccupied by Shinji, 1940a: 162.
Distribution: Japan (Hokkaido, Honshu, Shikoku, Kyushu).
Proanoplomus longimaculatus
Proanoplomus longimaculatus Hardy, 1973: 268. HT F (UZMH), Burma. Kachin: Kambaiti [25°24'N
98°9'E], 2000 m. Depository misstated?, NRS?
Distribution: Burma.
Proanoplomus nigroscutellatus
Proanoplomus nigroscutellatus Zia, 1964: 45. HT M (IZAS), China. Yunnan: Shishong-Baanna
[Xishuangbanna], 1200 m. Latest reference: Hancock, 1999: 938.
Distribution: e. India, China (Yunnan).
Proanoplomus omeiensis
Proanoplomus omeiensis Zia, 1964: 47. HT F (IZAS), China. Sichuan: Omeishan [Emei Shan].
Distribution: China (Sichuan).
Proanoplomus spenceri
Proanoplomus spenceri Hardy, 1973: 273. HT F (BBM), Vietnam. Fyan, 1200 m.
Distribution: Vietnam.
Proanoplomus yunnanensis
Proanoplomus yunnanensis Zia, 1964: 46. HT F (IZAS), China. Yunnan: Shishong-Baanna
[Xishuangbanna]. Latest reference: Hancock, 1999: 938.
Proanoplomus trimaculatus Hardy, 1973: 274. HT M (UZMH), Laos. Nam Tiene. Distribution: Laos.
Distribution: e. India, China (Yunnan, Guangxi), Burma, Thailand, Laos, Indonesia (Java).
189
BIOTAXONOMY OF TEPHRITOIDEA
23. RHAIBOPHLEPS
Rhaibophleps Hardy, 1973: 203. Type species: seclusa Hardy (OD). Latest reference: Hancock, 1999:
938.
Rhaibophleps seclusa
Rhaibophleps seclusa Hardy, 1973: 204. HT M (BBM), Thailand. Nakhon Ratchasima: “Khorat
Prov.”, 300 km NE. Bangkok, Sakaerat [Ban Huai Sakae Rat], 300-400 m.
Distribution: Thailand, Laos, Cambodia.
24. SINANOPLOMUS
Sinanoplomus Zia, 1955: 64. Type species: sinensis Zia (OD). Latest reference: Hancock, 1999: 938.
Sinanoplomus fasciatus
Urophora fasciata Walker, 1856: 134. LT F (BMNH), Malaysia. Sarawak. Lectotype designated by
inference of holotype by Hardy (1959: 226). Latest reference: Hancock, 1999: 938.
Carpophthoromyia borneensis Hering, 1952: 283. HT F (RNH), e. Borneo.
Distribution: Malaysia (Sarawak, Sabah), Indonesia (Kalimantan).
Sinanoplomus sinensis
Sinanoplomus sinensis Zia, 1955: 64. HT F (IZAS), China. Guangdong: Lianxian. Type data: Wang,
1998: 22.
Distribution: China (Guangdong).
25. SPILOCOSMIA
Spilocosmia Bezzi, 1914: 327. Type species: bakeri Bezzi (OD). Latest reference: Hancock and Drew,
1999: 758.
Prospilocosmia Shiraki, 1933: 212. Type species: punctata Shiraki (= bakeri Bezzi) (MO). Proposed
as a subgenus of Spilocosmia.
Spilocosmia bakeri
Spilocosmia bakeri Bezzi, 1914: 327. ST M (Baker), Philippines. Luzon, Laguna: Mt. Makiling.
Lectotype designation of Hardy (1969: 481) invalid; syntypes currently in MCSNM. Latest references:
Wang, 1998: 45; Hancock and Drew, 1999: 759.
Prospilocosmia octavia Munro, 1935b: 256. HT M (DEI), Taiwan. Taihoku. Synonymy: Hancock and
Drew, 1999: 759.
Spilocosmia (Prospilocosmia) punctata Shiraki, 1933: 214. ST B (NTU), Taiwan. Musha; Arisan;
Rikiriki; Taito; and Urai. Synonymy: Hancock and Drew, 1999: 759.
Prospilocosmia punctata f. kotoshoensis Shiraki, 1933: 216. ST B (NTU), Taiwan. Kotosho.
Synonymy: Hancock and Drew, 1999: 759.
Spilocosmia incompleta Wang, 1998: 45. HT F (CAS), China. Zhejiang: Moganshan (30.6°N,
119.8°E). Synonymy: Hancock and Drew, 1999: 774.
Spilocosmia octavia: Hardy, 1988: 115; Wang, 1998: 45.
Distribution: China (Zhejiang), Laos, Vietnam, Japan (Ryukyu Is.), Taiwan, Philippines (Luzon, Leyte,
Mindanao), Indonesia (Nusa Tenggara).
26. TAENIOSTOLA
Taeniostola Bezzi, 1913: 119. Type species: vittigera Bezzi (OD). Latest reference: Hancock and
Drew, 1999: 761.
Taeniostola striatipennis
Taeniostola striatipennis Hering, 1941b: 68. HT M (MNM), Malaysia. Sabah: Kinabalu. Latest
reference: Hancock and Drew, 1999: 761.
Distribution: Malaysia (Sabah).
190
Kovac et al. / Catalog and classification of Gastrozonini
Taeniostola vittigera
Taeniostola vittigera Bezzi, 1913: 119. ST B (ZSI), Bangladesh. Sylhet; and India. Assam: Lungleh.
Latest reference: Hancock and Drew, 1999: 762.
Taeniostola connecta Hendel, 1915: 436. ST M (MNM), Taiwan. Kosempo. Synonymy: Hancock and
Drew, 1999: 762.
Taeniostola apicata Hering, 1938a: 250. HT F (ZSZMH), Indonesia. Kalimantan: Bidang Menabai,
700 m. Synonymy and type data: Hancock and Drew, 1999: 762; holotype probably destroyed.
Taeniostola plagiata Hering, 1938a: 245, Incosp. apicata Hering. Hardy 1973: 210 (first revisor).
Distribution: Bangladesh, India (Assam, Mizoram), China (Yunnan), Burma, Thailand, Laos, Taiwan,
Malaysia (w., Sarawak, Sabah), Indonesia (Kalimantan).
27. XANTHORRACHIS
Xanthorrachis Bezzi, 1913: 137. Type species: annandalei Bezzi (OD). Latest reference: Hancock
and Drew, 1999: 764.
Xanthorrachis annandalei
Xanthorrachis annandalei Bezzi, 1913: 138. HT F (ZSI), Burma. Karen: Dawna Hills, 2000-3000 ft.
Latest references: Wang, 1998: 48; Hancock and Drew, 1999: 765.
Carpophthorella scutellomaculata Hering, 1951: 7. HT F (BMNH), India. Kerala: Anamalai Hills,
4000-5000 ft. Synonymy: Hardy, 1988: 119.
Distribution: India, China (Yunnan), Burma, Thailand, Laos, Vietnam, Indonesia (Java).
Xanthorrachis assamensis
Xanthorrachis assamensis Hardy, 1973: 283. HT M (BMNH), India. Assam: N. Khasi Hills, lower
ranges. Latest references: Wang, 1998: 48; Hancock and Drew, 1999: 766.
Distribution: India (Assam, Arunachal Pradesh), China (Xizang, Yunnan).
Xanthorrachis sabahensis
Xanthorrachis sabahensis Hardy, 1988: 119. HT M (BBM), Malaysia. Sabah: Sandakan Bay (NW.),
Sepilok Forest Reserve, 1-10 m. Latest reference: Hancock and Drew, 1999: 766.
Distribution: Malaysia (Sabah).
EXPLANATIONS TO TAXONOMIC CHANGES MADE IN THE CATALOG
As mentioned earlier, the catalog and key presented here are neither the result of a
comprehensive revision nor a cladistic analysis. Therefore, we have restricted changes to the
following:
1. Ceratitoides nigromaculata Hendel was separated from other congeners listed by Hancock
(1999), based on a study of specimens collected by us in Cameroon. Males have more or less
equally long abdominal tergites, unlike the enlarged second segment shown in Hancock’s
figure (Fig. 15) which, together with wing pattern, appears to be the main character used to
maintain Asian species in this genus. With one exception, these Asian species are either
newly placed in Pardalaspinus (Acroceratitis aberrata Hardy, A. adnata Hardy,
Chelyophora maai Chen, Ceratitoides namtamai Hancock, and C. sikhimensis Hancock, all
new combinations), or are returned to this genus. Carpophthoromyia cinereofasciata
Meijere is moved from Ceratitoides and placed in Proanoplomus (n. comb.), which it seems
to fit better based on wing pattern characters (see couplet 17 of key to genera above);
2. The wing figure of Acrotaeniostola morosa Hering by Hancock and Drew (1999, Fig. 16;
bands narrow, pale, with wide spaces between them) does not conform to that by Hering
(1938, Fig. 13; bands wide, dark, with narrow spaces in between) but does match Wang’s
191
BIOTAXONOMY OF TEPHRITOIDEA
(1998) figure and description of A. longicauda Wang. Hence, the latter species is reinstated
from the synonymy proposed by Hancock and Drew (1999).
ACKNOWLEDGMENTS
We are grateful to Dr. Stuart McKamey, Dr. Masahiro Sueyoshi (USNM) and Mrs. Irina
Zonstein (TAU) for critically reading the manuscript. Irina also tested the key against
specimens. Thanks are due to Dr. Rosli Bin Hashim (University of Malaya), who is in charge of
the Ulu Gombak Field Studies Centre (UFSC), where DK and PD were stationed during their
research work in Malaysia and to Decha Wiwatwitaya from the Kasetsart University, Bangkok,
for help during the field work in Thailand (DK). We also wish to acknowledge the help of the
UFSC staff, especially Bah Tera, the main warden. DK, AF and PD wish to thank the DFG
(“Deutsche Forschungsgemeinschaft”) for funding their work in Malaysia and Thailand for two
years.
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