tests nice

a mee
Pisses tee

3 0

test eae
sprig tie
ait

a
Ms aR wah: aN y's ¥
‘es a chMaN) ati

<

iy See"

¥ ; a 1y
Pai ee

: ‘ '
mi,

oe

1\f

LR ae

Li 2 ee BCR

pas (Ste HAS)
BAY

Bee, ony :

: t é ‘ ia t rf - ee
iar x 7% 7

=
ya i
—_—
; 7%
,
=
9
»

\

d <q \

BANS i/o
Le fe

A REVISIONARY Se Silay
OF THE RUTILIINI (DIPTERA:
TACHINIDAE), WITH KEYS TO

THE DESCRIBED SPECIES

R. W. CROSSKEY

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 19
LONDON : 1973

—

“ae
———

Ce ae
aerate

a
hn

o\

(S* =e
A REVISIONARY CLASSIFICATION OF THE
RUTILIINI (DIPTERA : TACHINIDAE), WITH

KEYS TO THE DESCRIBED SPECIES

NI

oy

J

2 8 MAR {9

Po.
V4 as

BY

ROGER WARD CROSSKEY

Pp 1-167; 109 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY

Supplement 19
LONDON : 1973

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY) instituted in 1949, 1s
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
veady. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Supplement 19 of the Entomological
series. The abbreviated titles of periodicals cited follow
those of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. nat. Hist. (Ent.). Suppl.

© Trustees of the British Museum (Natural History), 1973

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 22 February, 1973 Price £6.50

A REVISIONARY CLASSIFICATION OF THE
RUTILIINI (DIPTERA : TACHINIDAE), WITH KEYS
TO THE DESCRIGED SPECIES

By R. W. CROSSKEY

CONTENTS

Page
SYNOPSIS 3
INTRODUCTION 2 , 4
MATERIAL AND METHODS . ; : : ; : GI
ADULT CHARACTERS AND THEIR TAXONOMIC “VALUE 8

Kery To TRIBES OF THE SUBFAMILY PROSENINAE IN THE ORIENTAL AND
AUSTRALASIAN REGIONS : : ; F : F 17
TRIBE RUTILIINI BRAUER & BERGENSTAMM. ; : : : : 18
Key to the genera of Rutiliini.. : - : : , ‘ 2:
Genus Formosia Guérin-Méneville : : é : . P 2°
Genus Formodexia gen. n. . , . ‘ : : : : 38
Genus Rutilodexia Townsend , F : : : : f 40
Genus Rutilia Robineau-Desvoidy : Z : ; : : 42
Genus Amphibolia Macquart : : : : ; : 93
Genus Chrysopasta Brauer & Bergenstamm : : 3 - 2 102
Genus Prodiaphania Townsend . ® Z : : : : 107
Genus Chetogastey Macquart . : F . ; ‘ : 112
LrEcToTyPE DESIGNATIONS : : : . : ; : E 116
NEOTYPE DESIGNATIONS . : ‘ : 121
SUMMARY OF THE PROPOSED CLASSIFICATION OF THE Ruriwunt ; : 127
INDEX-CATALOGUE OF SPECIES-GROUP NAMES AND THEIR TYPES ; : 130
ACKNOWLEDGEMENTS ‘ P ; ‘ ‘ : : P ‘ 140
REFERENCES . : : : F : ; : ’ : : 140
ILLUSTRATIONS ; : : : ; : , : - 144
INDEX TO GENUS-GROUP NAMES . : : : : . ; ‘ 164
INDEX TO SPECIES-GROUP NAMES : 4 : ; : : F 165

SYNOPSIS

A comprehensive re-classification is proposed for the Rutiliini, a Tachinid tribe confined to the
Oriental and Australasian Regions. Diagnoses and identification keys are provided for all
genera and subgenera recognized, and all nominal species are appropriately placed in the
classification after examination of the types. Keys are given, under their respective genus-group
taxa, to almost all of the 121 described species that are recognized as valid; it is emphasized
that the species keys are only tentative, as species limits are often uncertain and several un-
described species are known. ‘The proposed classification is summarized and an index-cata-
logue is provided to all nominal species-group taxa and their primary types. One new genus,
one new subgenus and seven new species are described; 47 specific names and 15 genus-group
names are newly placed in synonymy; 29 lectotypes and 12 neotypes are newly designated;
and one new name is proposed for a preoccupied secondary homonym. There are 26 new
combinations (excluding those implied by new synonymy).

4 R. W. CROSSKEY

INTRODUCTION

THE Rutiliini include the largest and most handsome flies to be found in the Tachini-
dae, and the brilliant metallic colours and conspicuous patterns of many species
make them outstandingly attractive insects. The group is not well known outside
of Australia, where it pre-eminently occurs, and the glinting beauty of the metallic
forms comes as something of a surprise to the non-specialist accustomed to think of
the Tachinidae as just about the dullest of the Diptera.

Because of their showy colouring and flower-loving disposition the Rutiliines were
among the first Diptera to be collected in Australia, and most of the expeditions of
the late 18th and early 19th centuries that had touched at ‘New Holland’ (Australia),
“Van Diemen’s Land’ (Tasmania) or the islands of Melanesia brought back specimens
that were described by the European naturalists. The most venerable surviving
example is Fabricius’ type of Rutilia retusa in the Banks collection in London which
was described in 1775 and was probably collected on one of Captain Cook’s voyages.

From the writings of the earliest workers on the group it is clear that they were
much impressed by the wonderful ‘new’ flies coming from Australasia, and this is
attested by the names—such as imperialis, mirabilis, regalis, splendida—they bestow-
ed on them. Some of the first describers were not primarily dipterists, and the best
early work on Rutiliini was produced by the coleopterist Guérin-Méneville (1843).
By 1850 the rather surprising number of 44 nominal species had been described, and
a further 40 nominal species were named before the end of the 19th century (mainly
by Macquart, Walker and Bigot)—many of the names referring to features of the
colour and marking.

The Rutiliini are more difficult to classify satisfactorily than would be supposed
from their obvious appearance, and the fact that the bright colours and bold patterns
can be relatively easily described has had one definite disadvantage in the systematic
history of the group: it has tempted later workers to think that they could recognize
the species described by their predecessors on the basis of the descriptions, without
recourse to the types. Often this was not so, and both generic and specific nomen-
clature have been much confounded by the misapplication of names. The Rutiliini
has also had its share of ‘splitters’, notably Townsend and Enderlein, creating what
Paramonov (1968 : 351) amusingly called their ‘deluge’ of new genera. The mis-
identifications and generic splitting, together with the fact that no previous compre-
hensive study of old types was possible, has long meant that the systematics of
Rutiliini have been in a bad state—as Paramonov put it we are ‘left with the task
of solving the riddles’. He himself had made an excellent start on this task, and
Paramonov’s (1968) posthumously published paper dealing with the genera other
than Formosia and Rutilia is the best work on Rutiliini that has up to now appeared
—in a class vastly superior to the superficial splitter’s nightmare produced by
Enderlein (1936) as a ‘Klassifikation der Rutiliinen’, which Paramonov so rightly
deplored. Paramonov did not live to complete his projected work, and there was
therefore no revision up to now of Formosia s.l. and Rutilia s.l1., which together
constitute about 70% of the Rutiliini. -

The present paper is an attempt at a new comprehensive classification of the whole
tribe. The work on which it is based grew gradually and inevitably out of the much

REVISIONARY CLASSIFICATION OF RUTILIINI 5

simpler objective of preparing an up-to-date catalogue of the Australian Tachinidae,
a task which sounded easy enough but soon proved to be impossible without a really
thorough revision of the Rutiliini. This group forms a very dominant element in
the Australian Tachinid fauna, including at present about a quarter of the known
species, and a catalogue of this large group based only on the muddled literature
would have been not only useless but positively misleading. A dependable cata-
logue had to be based on a study of all the types, especially those of the early
authors whose nominal species had been neglected or misinterpreted, so that realistic
generic assignments and at least the obvious synonymies in genus-group and
species-group names could be worked out. But even to make a generic assignment
of a nominal species presupposes some concept of the generic limits, and when
trying to place species into genera it soon became obvious that none of the existing
generic classifications (Townsend, 1938; Enderlein, 1936; Paramonov, 1968) could
be used satisfactorily, although that of Paramonov was outstandingly superior to
any other (in fact my own classification here presented is in close accord with that
of Paramonov in the number and scope of recognized genera). In short, the only
way in which the Rutiliini could be satisfactorily catalogued was by first preparing a
full-scale revisionary classification of the whole tribe.

The classification proposed here is based on the ‘old-fashioned’ methods of
orthodox taxonomy, but the Rutiliini is a group which might lend itself well to the
computer techniques of numerical taxonomy.. The use of mathematical methods
was considered at one stage of the work but it was decided to abide by conventional
methods for the time being, in the hope that later on an Australian student might
take up the group and test it by mathematical methods on far more material than
is currently available. Two of the main difficulties with classical taxonomy in the
Rutiliini are those of ranking of segregates and delimitation of species. Broadly
speaking it is easy enough with suites of characters taken in combination to define
segregates within the tribe (or, looked at the other way, aggregates of species) but
it is a very subjective matter whether these are ranked as species-groups or genus-
group categories; I have not always been completely happy at the choice of rank,
for it is difficult to ‘balance’ recognizable segregates, but I am convinced that the
groupings I recognize are natural entities whatever rank they may be accorded.
On what is a species I am less happy, as there are real difficulties in several genera
and subgenera in determining specific limits (some entities that are apparently
species have distinctive and constant male genitalia, for example, but others either
show no genitalic differences or some baffling variation).

Two particular aspects of the work require comment, the examinations of old
types and the male genitalia. I have been able to see very nearly all of the types
of the early authors (Bigot, Erichson, Fabricius, Gerstaecker, Guérin-Méneville,
Macquart, Walker, Wiedemann), only very few of which are lost; the types of
Donovan, Gray and Robineau-Desvoidy are all lost. Examinations of these old
types have enabled many formerly enigmatic names to be placed, either as valid
names or as synonyms (because of the difficulty in some groups of deciding on specific
limits, synonyms have only been established if there is very perfect agreement of
types), and have unmasked several misidentifications.

6 Ro W.. CROSSE Y

A particular study has been made of the male genitalia. Malloch (1929, 1936),
Engel (1925) and Paramonov (1968) published a few figures of male terminalia,
but made no systematic study of them. In the present work the male genitalia
have been examined for the great majority of species in order to assess their useful-
ness in classification; the outcome has been to find that they have very limited use
for supraspecific classification but in many groups provide valuable characters at
specific level (for further detail see the section on taxonomic characters).

The larger museum collections of Rutiliini, especially those at Canberra and
London, contain specimens of several species that are obviously undescribed. I
have not attempted to place these in the keys to species, which have been drawn
up to cover only those species already described (as the title of this work states),
but I have here described seven new species and these are placed in the keys. The
new species have been described here for definite purposes: either because they
show characters that significantly extend the range of form or colour previously
known in the taxa to which they belong, or because they extend the previously
known geographical range of their genus or subgenus, or to clarify species-complexes
in which there was no available name for one of the constituents. Other new species
known in collections have not been described because there are no cogent reasons
at present why they need be named.

I must advert in this Introduction to the classification, briefly referred to above,
of that entomological Jack-of-all-trades, Giinther Enderlein. The paper of Enderlein
(1936) on the Rutiliini was, luckily, his only venture into the Australian Tachinidae,
except for his description of Microtropesa violacescens. It contains the usual lavish
erection of unnecessary Enderleinian genera, 16 new genera in all, of which only one
is given any recognition in my classification (this as a subgenus of Rutilia); I agree
whole-heartedly with the late Dr Paramonov’s stricture that ‘these genera are
unwarranted and very often misleading, as they are based on erroneously identified
species of the early workers’. It should be added, however, that many of Enderlein’s
species are valid, even if their descriptions are excessively perfunctory by 2oth
century standards.

The genera Oxyrutilia Townsend, Ola Paramonov and Ruya Paramonov do not
appear in the present work although they were placed by their describers in the
Rutiliini: the first of these genera is a synonym of Nemoraea Robineau-Desvoidy
(tribe Nemoraeini) as Paramonov (1968 : 351) established, and the other two are
excluded from the Rutiliini as here defined for the reasons given on p. 22.

The figures accompanying the text of this paper have all been drawn personally.
Those of the male genitalia are in the form of simple outlines with the hair vestiture
omitted, as it is shape which is important for recognition and highlights the essential
differences (or resemblances)—over-fussy drawings with every hair in place are
usually not a virtue in Calyptrate taxonomy (unless of course the characters, as
sometimes happens, reside in the vestiture itself). Some of the drawings are semi-
schematic, notably those of chaetotaxy, which is often better represented by the
pores than by the bristles themselves.

Finally in this Introduction I must refer to my use of the subfamily name Pro-
seninae. For some years this name has been current for the very large subfamily

REVISIONARY CLASSIFICATION OF RUTILIINI 7

once known (wrongly from the nomenclatural viewpoint) as the Dexiinae, and I
have here continued to use the subfamily name Proseninae although it is clearly not
the oldest available name for the taxon. The family-group name Proseninae dates
from 1892, and is pre-dated by several family-group names proposed by Brauer &
Bergenstamm (of which Rutiliidae itself is one) and probably by other even earlier
names. It will be a very complex nomenclatural question to resolve which of the
many family-group names in the Tachinidae are based upon genera belonging in
the “Proseninae’, and which of them should be brought into use as the valid name
for this subfamily; until this can be done thoroughly, with a real prospect of long-
term stability, I consider it best to continue using the name Proseninae as I have
done in the present work.

MATERIAL AND METHODS

The revisionary work presented here has been based on the extensive collection
of Rutiliini housed at the British Museum (Natural History) and on the assembly
of as many of the primary types as possible from overseas museums. Most of the
types in depositories in Sydney and Canberra were examined during a visit to
Australia in 1965, and Macquart’s types (which could not be borrowed) were studied
at the Paris museum in 1969. Primary types have been examined of 151 species-
group nominal taxa out of the 168 for which type-material is known to be still in
existence; for the remaining few names type examination was either not essential
because the species were recently and very well described, or was not possible
because types could not be loaned and there was no opportunity to examine them
(e.g. Macquart’s types of Amphibolia valentina and Diaphania testacea in Lille).
Some types are lost (surprisingly few in view of the relative antiquity of many of
the names) and a few have not been located but may still exist. The primary types
of the 168 nominal taxa with located types are distributed as follows: 54 in British
Museum (Natural History), London; 49 in Australian collections (Australian National
Insect Collection, Canberra; Australian Museum, Sydney; School of Public Health
and Tropical Medicine, Sydney); 34 in Museum fiir Naturkunde der Humboldt-
Universitat, Berlin; 14 in Museum National d’Histoire Naturelle, Paris; 12 in small
European museums (Brussels, Leiden, Lille, Stockholm and Vienna); and 5 in North
America (New York and Washington). Seven new species are described in the
present paper from material in the British Museum (Natural History) and their
holotypes are in this museum.

Small collections of Rutiliini from the museums at Oxford, Leiden and Paris, and
from my own collecting in New Guinea and New Britain, were at hand and taken into
account during the preparation of this paper.

The early stages of Rutiliines remain almost completely unknown, and only adult
flies have been studied. These do not require any special techniques, but some
comment may be helpful on preparation and figuring of male genitalia. Rutiliini,
being bulky flies, have sizeable male genitalia which are difficult to slide mount
satisfactorily, even with cavity slides, though permanent slide preparations should
always be made using such slides; nearly always the hypopygium rolls at least slightly

8 R. W. CROSSKEY

out of position in a permanent preparation so that an undistorted profile or apical
view is not given. It is therefore vital to examine the hypopygium in fluid and to
compare the shapes of the surstyli and cerci against the figures before slide mounting.
All the accompanying figures of the male epandrium, surstyli and cerci have been
drawn from whole hypopygia removed from the flies, slightly softened in potassium
hydroxide solution, and mounted in fluid to show either the shape in profile (lateral
view) or in apical view. Some of the specific differences in shapes of surstyli and
cerci are very subtle, and to get comparable posterior views the hypopygium has
always been orientated in such a way that the sclerotized dorsal part of the epan-
drium is just visible above the ‘open’ membranous part. This should be kept in
mind when comparing an apical view of the male hypopygium against the figures.
As a rule it is not possible to obtain very satisfactory views of the male hypopygium
by extracting it in situ on relaxed flies; some very important features lie at the
bases of the surstyli and may be concealed under the epandrium and overlooked
unless the hypopygium is completely removed, so that it can be viewed from various
angles in fluid at least while an identification is being made. (Examination of the
male hypopygium is only needed at the specific level, and as species do not normally
show important characters on T5 or on sternite 5 it does not usually matter greatly
if the surrounding parts of the abdomen are slightly damaged when the hypopygium
is removed.)

When describing the leg chaetotaxy the convention is followed of imagining the
leg to be extended at right-angles to the longitudinal axis of the fly, when: ad =
anterodorsal, av = anteroventral, pd = posterodorsal, and pv = posteroventral.
The close-set comb of setulae on the hind tibia of many forms is referred to as the ad
fringe, using the conventional positional terminology ‘ad’; this equates with Para-
monov’s (1968) term ‘externodorsal’.

The abbreviations used for thoracic chaetotaxy are: acr, acrostichal setae; dc,
dorsocentral setae; za, intra-alar setae; ph, posthumeral setae; stpl, sternopleural
setae. Position before or behind the transverse suture of the mesonotum is indicated
by prst (presutural) and post (postsutural) respectively, in the normal convention;
hence prst dc indicates presutural dorsocentral setae, post 1a indicates postsutural
intra-alar setae.

Abdominal tergites are indicated by the letter T followed by the appropriate
number; the composite first apparent tergite is Tr + 2, the last visible tergite T5
(Text-fig. 28). (It should be noted here that Paramonov (1968) referred to abdominal
tergites by their apparent number, not by their correct numbering on the basis of
morphological segmentation: his ‘first’ tergite is correctly Tx + 2, his ‘second’ and
‘third’ tergites are T3 and T4, and his ‘fourth’ tergite is T5.) For the convenience
of Australian workers the terminology of the parts of the male hypopygium is that
adopted by Colless & McAlpine (1970) in their work on Australian Diptera.

ADULT CHARACTERS AND THEIR TAXONOMIC VALUE

At present only morphological characters of adult flies are available for classifica-
tion and species recognition. The following account details the characters that

REVISIONARY CLASSIFICATION OF RUTILIINI 9

have use in taxonomy at supraspecific and specific levels, and indicates briefly some
of the features which (at any rate on present evidence) have no taxonomic usefulness.

BopDy COLOUR AND POLLINOSITY

These are so closely interlinked, the appearance and pattern being largely deter-
mined by the distribution of the pollinosity and its density, that they are considered
together. Useful specific characters, and to some extent supraspecific characters,
are provided by the extent of visible metallic coloration ; some taxa are characterized
by having the parafrontals (sometimes also the parafacials) brilliantly metallic
instead of pollinose (as is usual), and some have the genal dilations and epistome
partially or entirely metallic. Metallic colouring of the abdomen is especially
important, several segregates being characterized by the occurrence of transverse
metallic bands or spots on most of the tergites (usually the metallic colour being
golden green to coppery red). The presence and extent of metallic coloration on
the abdomen provide useful characters at several levels, but in some groups there is
conspicuous intraspecific variability; in some Chrysorutilia species, for example,
the metallic pattern may consist of continuous transverse bands or of broken or
partially coalesced metallic spots in the same species (and there is a tendency for
females to have more complete banding than the males, which more frequently
have the metallic pattern is discretely isolated spots).

The ground colour of the head and its overlying pollinosity often provide useful
characters for distinguishing allied species, and some supraspecific aggregates may
have a particular head colour (a brilliant golden yellow head is a common form
which appears in unrelated groups). The thorax normally shows traces of whitish
pollinosity over the prescutum at least, and conspicuous spots of ‘thick’ white
pollinosity occur in many forms in a standard pattern on the thoracic dorsum and
often on the mesopleura and sternopleura; the presence of such bold spots is often a
specific or group character. In some species the spots, especially those on the
mesopleura, may have a more ‘shifting’ appearance with the direction of the light
than in others. The mesonotum typically shows four blackish vittae, and the bold-
ness and extent of interruption of the vittae at the transverse suture sometimes
provide (somewhat tenuous) specific differences. Some species and groups are
characterized by having the thorax and abdomen uniformly dark, in which case
mesonotal vittae are not evident.

Leg colour and, to a minor extent, antennal colour can provide useful specific
characters. In some forms, however, such as Chrysopasta there appears to be intra-
specific variability in leg colour, and it appears likely (but is yet to be proven)
that some species of Microrutilia may have sexually dimorphic leg colour (black in
males, reddish yellow in females).

Hair colour is of no importance at the supraspecific level and its significance at the
species level is far from clear. Several very closely related and virtually inseparable
species differ by having either black or mainly yellow pleural hair on the thorax,
and it is possible that some species are polymorphic in hair colour; it seems probable,
too, that some species may be sexually dimorphic in this feature, males having black
pleural hair and females having yellow pleural hair (as in some species of the Goniine

10 R. W. CROSSKE ¥

genus Winthemia Robineau-Desvoidy). (In the present work it has been impossible
to come to any definite conclusion about polymorphism or sexual dimorphism in
hair colour, and nominal species have been treated as valid if their types differ in the
colour of the pleural hair.) Colour of the parafrontal, genal, postbuccal, coxal,
femoral, tibial, and ventral and apical abdominal hair has limited value as a specific
character. Bristles of the chaetotaxy are nearly always black, but some of the
bristling in a few species of Chrysorutilia is golden red (especially the postalar and
vertical setae and the postocular setulae); the significance of this abnormal bristle
colour is not clear.

CHAETOTAXY AND HAIRING

Chaetotaxy in general. The most striking feature of the chaetotaxy in the
Rutiliini is its instability, much of the bristling being less constant in this tribe than
in other Proseninae and incomparably less constant than in the higher Tachinidae
(Tachininae and Goniinae). In the Goniinae, particularly, the principal bristles are
very constant in arrangement and size, and whole tribes may have a uniform arrange-
ment of many of the setae—for example all the Sturmiini have 3 + 4 dorsocentral
setae. In the Rutiliini there is not only intraspecific variability among many of the
setae in their number, but also in their degree of development and differentiation
from the surrounding hair; there is often also a lack of bilateral symmetry in the
bristling of individual specimens, and a few species are sexually dimorphic in certain
of the setae (e.g. median marginal setae present on abdominal T3 in females but
absent in males). Failure to appreciate the inconstancy of the chaetotaxy led
Enderlein (1936), working with rather limited material, to erect several untenable
genera on the basis of supposed chaetotactic differences, and in the case of Hega
Enderlein (based on male specimens) and Chromocharis Enderlein (based on females)
to propose genera for the opposite sexes of the same taxa.

In spite of the variability in the chaetotaxy, and the fact that almost any chaeto-
tactic character will fail in the occasional specimen, it is none the less the case that
the chaetotaxy provides some of the most important characters for supraspecific
classification—and that several of them are essential key characters by which genera
and subgenera can be readily enough distinguished. The existence, however, of
isolated specimens showing chaetotactic characteristics which conflict with the
norm for their nominal taxon has always to be kept in mind (especially as it is
virtually impossible to cover every conceivable variant in odd specimens in a
practical key).

Head chaetotaxy. There are few features of the head setae of use in classification.
The ocellar setae are normally small or virtually absent and but little differentiated
from the long hair of the ocellar triangle. The vertical setae are represented only by
the inner pair, which shows no useful features. The frontal setae are usually very
small and fine (Text-fig. 3), often little more than short hairs, and frequently the
two rows of frontals do not meet at the tips; critical examination might yield minor
specific differences, but there are no obvious taxonomic characters in the frontal
setae. Proclinate orbital setae are always absent in males and are often extra-

REVISIONARY CLASSIFICATION OF RUTILIINI IL

ordinarily reduced or absent in females; when present in females their size and num-
ber often vary, and may be different on each side of the same specimen, but never-
theless there is a tendency for their normal presence or absence in different segre-
gates. The vibyissae are sometimes long and strong but most often are weakly
differentiated from the peristomal setae and from the small setulae above the main
vibrissae (Text-fig. 3); they have no real taxonomic use. The facial ridges are bare,
but in Chrysopasta small setulae extend up them further from the main vibrissae
than is usual. The row of postocular setulae varies somewhat in length and might,
if sufficiently studied, show minor specific differences in males.

Thoracic chaetotaxy (Text-fig. 9). The most valuable chaetotactic characters are
on the thorax, and several groups of thoracic setae provide generic and subgeneric
characters. The basic number of humeral setae is four, two on the outer half and
two on the inner half of the humeral callus, but in some Rutilia s.l. the inner pair is
absent or virtually-so. The posthumeral setae (pf) are typically developed in some
genus-group segregates and undeveloped in others but are of very minor taxonomic
usefulness; often their development is variable, and in males ph setae may be un-
differentiated from the prescutal hair although distinct in females. The acy and dc
setae are especially variable and almost no use has been made of them in the present
work; in Formosia the presence of a distinct prst dc seta in one group and not in
others has been noted. Some segregates characteristically show some strong slightly
spiniform setae on the scutum between the hindmost setae of the acy and dc rows
(so that the scutum shows a rather continuous transverse row of strong setae imme-
diately before the scutellum); the presence or absence of these supernumerary
prescutellay setae provides a minor taxonomic character. The pre-alar and supra-
alar setae are extremely variable in development and have no taxonomic value;
the pra seta may be present or absent in the same species or on the two sides of the
same specimen, and there may be several supra-alar setae or only one in the same
species. The notopleural setae are normal (1 + I) in almost all forms, but a few
curious species from the Philippines have a well developed third notopleural seta
standing very close to the normal posterior notopleural (i.e. I + 2 notopleurals),
the hind pair standing on an unusually prominent knob-like swelling of the noto-
pleuron. Intra-alar setae occur only as post 1a setae on the scutum, and there is
never a prst ia seta in Rutiliini, but the post za are often intraspecifically variable;
typically in any species there may be only one post ia, but almost always occasional
specimens will be found in which a second or even third fost ia is present, and in
species in which there is normally more than one there may sometimes be found
only a single post ia; consequently the za setae have very limited taxonomic value.

The most dependable thoracic setae for taxonomic purposes are the postalar setae
on the postalar callus, the sternopleural setae, and the scutellar setae. Omitting
Chetogaster (which has two postalars like normal Tachinidae) the Rutiliini are very
exceptional in having more than the basic two setae developed on the postalar
callus, there being either three very strong setae or four or five (exceptional speci-
mens may even have six, with or without a weakly developed seventh seta). The
possession of either three postalars or more than three appears without doubt to
aggregate Rutiliine species into natural groupings, though the development of one

12 R. W. CROSSKEY

or more supernumerary postalar setae (in addition to the basic two) has evidently
occurred several times over in different evolutionary lines—hence different genera
and subgenera have either three or four (+) postalars, and the common possession of
either of these numbers does not indicate that the genus-group taxa involved are
necessarily phyletically close. Unfortunately the number of postalar setae is not
absolutely dependable in every specimen, for very rarely specimens do occur in which
the postalar callus has three setae on one side and four on the other (in this case it
seems from other characters that such specimens always, or nearly always, belong
to taxa in which 4+ is the norm for postalar setae). The sternopleural setae provide
some useful supporting characters in the definition of genus-group segregates, some
typically having only a single (posterior) stp] seta and others having either I + 1
or 2 + 1 stpl, but some variability occurs; for example groups normally having 2 + 1
may lack one of the anterior pair in some specimens or on one side of a specimen, and
groups normally having I + 1 stpl may have the anterior one virtually indistinguish-
able from the sternopleural hairing (especially in the male). No taxonomic charac-
ters have been discovered in the mesopleural, hypopleural, propleural or prostig-
matic setae; there is no definite pteropleural seta differentiated from the tuft of long
strong pteropleural hairing below the wing-base in any Rutiliini.

The scutellar setae show characters of value in the definition of genera and sub-
genera. The number and strength of marginal scutellar setae vary much, but the
position of the apical pair (whether inserted lower than or level with the other
marginals) provides a very dependable character (in all Formosia s.1. for example the
apical scutellars are level with the other marginal scutellar setae, whereas in Rutilia
s.l. they are set at an obviously lower level on the tip of the scutellum). In some
segregates the scutellum carries a transverse row of small but distinct preapical
setae lying in the same horizontal plane as the marginals, and these provide a useful
character (though there is some variability in their number and size it is nearly
always certain whether they can be classed as present or absent).

In some forms the thoracic setae (especially the supernumerary prescutellars, if
present, the scutellars, the postalars and some setae on the venter of the sternopleura
in front of the middle coxae) are markedly spiniform; in this case they are usually
stiffer, straighter and relatively shorter than in other forms. In some genus-group
segregates (notably Formosia s.l. and Rutilia s.str.) the development of strong
spiniform setae can be very striking, but it is difficult—because of intergradation
with forms with more normal bristling—to use the spiniform nature of the setae as a
taxonomic character.

Leg chaetotaxy. The legs provide few useful taxonomic characters. The setae
of the fore and mid legs (such as the pv setae of the fore tibia and ad setae of the
mid tibia) may vary in size and number and no useful features have been found.
Some forms have very heavy spiniform bristling on the mid and hind coxa but this
cannot be used as a dependable character. The development of the ad fringe of
the hind tibia, whether formed as a regular close-set comb or as a sparse irregular
row of ad setae and setulae, sometimes provides a helpful character, and is some-
times correlated with a different number of fd setae on the hind tibia (though this
can be intraspecifically variable or show sexual differences, e.g. pd setae absent in

REVISIONARY CLASSIFICATION OF RUTILIINI 13

male but present in female). In Formosia s.str. the whole anterior surface of the
fore coxa is haired, this feature distinguishing the segregate from all other Rutiliini
(in which the anterior surface is bare except near the apex). In the genus Prodia-
phania some valuable specific characters are provided by the setae of the ventral
surface of the hind femur, and in some species by a series of long setulae found on
the ad surface of the hind metatarsi of the male (such setulae occur nowhere else in
the Rutiliini).

Abdominal chaetotaxy. The presence or absence of long strong setae on T5, of
discal setae on the intermediate tergites, and of median marginal setae on T3 provide
useful characters at various taxonomic levels. The bristling of the inner ventral
ends of the tergites, where they nearly meet in the mid line of the abdominal venter,
varies much in strength and is sometimes spiniform; the direction of these setae,
whether pointing downwards (as in Formosia s.l.) or mainly backwards and only
slightly downwards (as in most other genera) has some value as a character at
genericlevel. The arrangement and nature of the vestiture of T5 are often especially
important, some segregates showing long haphazardly arranged setae, others one or
two regular transverse rows, and others having little more than sparse weak hairing;
some unusual species from the Philippines show short stubby setae irregularly in-
serted over most of T5. In many forms the abdominal chaetotaxy is exceptionally
strongly spiniform, but there are various degrees of ‘spiniformity’ in different genus-
group taxa and the spinous setae cannot in themselves be used as a taxonomic
character.

Hairing. The presence or absence, or extent, of hairing on different parts of the
body can provide valuable taxonomic characters at species-group, or genus-group
levels. In Rutilia s.1., for example, a valuable character for distinguishing the
subgenera Chrysorutilia and Ameniamima from other subgenera (and indeed from
all other Rutiliini) is the extent of hairing on the pteropleuron; in these segregates
the hairing on the pteropleuron extends well forwards on the sclerite, reaching to a
level much in front of the posterior stpl seta (Text-fig. 19), whereas in other Rutiliines
the anterior half of the pteropleuron is bare and there is virtually no hairing in front
of the level of the posterior stp/ seta (Text-fig. 20). In Formosia s.str. the whole
anterior surface of the fore coxa is haired, this feature distinguishing the segregate
from all the other Rutiliini (in which the anterior surface of the fore coxa is bare
except near the apex). Bare or haired parafacials distinguish many closely allied
species, and some genus-group segregates contain only species having fully haired
parafacials (e.g. Chrysorutilia). The barette is completely haired in nearly all
Rutiliines but some Chetogaster species have the hindmost part of the barette bare;
the type-species of Chetogaster and some closely allied species show a minute tuft of
fine hairs on the mediotergite beneath the lower calypter (infrasquamal setulae), but
| the mediotergite is totally bare in all other forms. Rutiliini normally have the
propleuron thickly haired, but very rare individual specimens (therefore no taxo-
nomic significance) have it bare.

The most important taxonomic characters of the hairing lie in the postalar wall
and the suprasquamal ridge (Text-figs 21-25), both of which may be haired (but

14 RK. We CROSSKEW

never both in the same taxon). In some genus-group segregates the postalar wall
(i.e. the vertical lateral declivity of the postalar callus, below the rounded margin
bearing the setae) has a thick tuft of long dense hair, but in most groups the postalar
wall is bare (or at most has just one or two hairs on its extreme upper part imme-
diately below the ridge of the callus) ; in other segregates the suprasquamal ridge is
most often haired, either with long dense bushy crinkled hair which is so thick that
the centre part of the suprasquamal ridge cannot be seen or with rather short sparse
hair under which the whole of the ridge is clearly visible (the nature of the hairing
therefore provides a useful character as well as its presence). A few segregates
(including the genus Rutilodexia, the subgenus Ameniamima and some species of
Rutilia s.str.) have both the suprasquamal ridge and the postalar wall bare, but most
often one or the other of these structures is hairy.

Presence or absence of hair on the prosternal membrane (Text-fig. 18) has some
taxonomic value. In Chrysorutilia and the aberrant species Rutilia micropalpis
there is hair on the anterior margin of the prosternum as well as on the membrane,
but hair actually on the prosternum itself does not occur in any other Rutiliini.

The hairing of the abdomen and male hypopygium shows no characteristics of
supraspecific taxonomic value, but there are sometimes minor differences at the
specific level in the length, strength and bushiness of the hairing, especially on the
epandrium and surstyli. Hairing on the surstyli provides some particularly good
specific characters in Prodiaphania.

Hairing on the arista ranges from extremely short micropubescence to moderately
long plumosity, and is generally similar in the species of any particular subgenus;
aristal hairing therefore provides a character of some minor value at the supra-
specific level.

Heap (Text-figs 1-3)

The most important taxonomic character at supraspecific level is the shape of the
facial carina, which Paramonov (1968 : 355) used in the first couplet of his key to
Rutiliine genera. Particular genera and subgenera usually show a moderately
constant facies in carina shape, but there is normally also some intraspecific and
interspecific variation and the character is not so easy to use in practice as Para-
monov’s key implies. In some segregates the carina forms a prominent convex
knob between the antennal bases and becomes slender and sharper towards the
epistome, while in others it is very broad along its length and has subparallel sides;
the latter form of carina often shows, superimposed upon its basic shape, either a
trace of a median sulcus or a median ridge. The efistome is always slightly promi-
nent in profile, and often subnasute, and is rather constant in shape in any segregate;
in forms with a bulbous facial carina the epistome is normally very prominent and
the face in profile is deeply concave between carina and epistome, but in forms with a
broad flattened carina the epistome is usually less prominent and not conspicuously
differentiated in profile from the epistome by a deep saddle. Most Rutiliines have a
well developed haired dilation on the gena (the so-called genal dilation), but in a few
forms (especially Rutilodexia) there is very little genal dilation and the dilated part

REVISIONARY CLASSIFICATION OF RUTILIINI 15

is widely separated from the vibrissal area (not reaching nearly as far forwards on
the head as the front of the eye, Text-fig. 5).
The eyes are well separated in both sexes in the majority of forms, but in Formosia

_s.l. many species have the male head nearly holoptic. In these the upper part of the

frons is almost obliterated, and the upper ends of the very attenuated parafrontals
may meet in the mid line (obliterating the upper part of the interfrontal
area completely). When the eyes are very nearly touching, the uppermost facets
are usually very conspicuously enlarged, and the closeness of the male eyes and
facet enlargement can provide useful specific characters. In all Rutiliini the eyes
are totally bare. The eye-height in relation to the width of the gena will probably
provide significant specific differences when sufficiently studied. The ocellar
triangle is exceptionally prominent in forms with the male head virtually holoptic
(a correlated feature).

The form of the buccal opening provides a character of some taxonomic impor-
tance. In most forms the opening at its narrowest (near the middle) is conspicuously
wider than the facial carina, but in the genera Prodiaphania and Formodexia the
buccal opening is unusually elongate and narrow (especially in the male) and at its
narrowest point is not or scarcely wider than the facial carina (Text-fig. 15). The
proboscis is of very uniform length, never greatly elongate, and the mentum has two
moderately distinct shapes (Text-figs 12 & 13) which are constant in any genus-group
segregate: in one shape the upper and lower edges of the mentum are subparallel
seen in profile so that the mentum is not noticeably tapering, and in the other the
upper and lower edges seen in profile distinctly converge apically so that the mentum
is tapering.

The antennae are always very small and their apices fall short of the epistomal
margin by a distance about equal to, or only a little less than, their own length; they
have no characters of supraspecific value, but the length of the third segment
relative to the second sometimes provides a useful specific character. The palpi
are minute (not longer than third antennal segment or basal thickness of the mentum)
in Prodiaphania but are well developed, long and slender, in other forms (a little
shorter than normal in Rutilia micropalpis and in Chrysopasta); they are sexually
dimorphic in Chetogaster (slender in males, spatulate or clubbed in females) but not
detectably so in other genera.

THORAX, LEGS AND WINGS

The structure of the thorax, legs and wings is extremely uniform and provides
very few taxonomic characters. In a few species the posterior part of the noto-
pleuron is produced as a knob-like swelling that is much more prominent than usual,
but otherwise the thoraces are alike throughout the tribe. Some forms have slightly
more elongate legs and tarsal claws than others, but not in any tangible way that
can provide taxonomic characters. The tegula (epaulet) has a pair or more of long
wiry setulae on its posterior edge in all Rutiliines except the one species of Formo-
dexia. In some forms, especially species of subgenus Donovanius, the wing mem-
brane is partially bare along some of the basal cells (at least no microtrichia visible
by entomological microscope, although the S.E.M. microscope might prove them to

16 R. W. CROSSKEY

be present in reduced form), and it appears that this might provide a useful specific
character. There are only minute differences in the relative proportions of the
costal sectors, but the degree of dilation of the costal base provides a character of
minor taxonomic use; some species have the costal base exceptionally strongly
flattened and widened (explanate), especially in the male, and the explanate costal
bases (i.e. the antecosta of Paramonov) then give the wings the appearance of having
basal ‘shoulders’ which are very easily visible to the naked eye (especially in flies
with the wings set back in the resting position). Partial or complete infuscation of
the wings occurs in some species, and can provide a helpful specific character.
Basicostal colour is also sometimes helpful.

ABDOMEN AND GENITALIA

Abdominal form. The abdomen is always very robust (Text-figs 28 & 29) and
often conspicuously broad and slightly flattened, and the shape is moderately
uniform in each genus-group segregate. The shape of T5—if convexly rounded and
tapering posteriorly or whether short, very broad and deeply hollowed medially—
provides an important character at supraspecific level. The degree to which the
sternites are exposed between the approximated ventral ends of the tergites does not
seem to provide any helpful characters except at the specific level in Prodiaphania.
The bilobed fifth sternite of the male shows little variation through the tribe as a
whole (Text-figs 30-35), the two lobes normally having a simple rounded shape
(Text-figs 30 & 31), but an unusual shape of fifth sternite occurs in Grapholostylum
(Text-fig. 33) and the lobes are straighter than usual apically in Microrutilia (Text-
fig. 32). In the subgenus Paramphibolia each lobe has a small blunt tooth or prong
on the inner edge towards the apex (Text-fig. 35), and this form of male fifth sternite
occurs nowhere else in the Rutiliini.

Male genitalia (Text-figs 36 & 39). The general structure of the male hypopygium
is exceedingly constant throughout the Rutiliini and there are really no characters of
importance at the supraspecific level; even at the specific level the taxonomic charac-
ters are almost confined to the surstyli, with a few useful characters in the cerci and
epandrium. The hypandrium (sternite 9), pregonites, postgonites and epiphallus —
(spinus) are virtually identical in every species, and the aedeagus itself has an
astonishingly constant form. The only difference observed in the aedeagus between
different species lies in the distiphallus, in which the relative lengths of the proximal
sclerotized part and the membranous distal part vary slightly. In the majority
of forms the membranous part of the distiphallus is about as long as the sclerotized
part or a little shorter (Text-fig. 37), but in a very few species (mainly the subgenus
Grapholostylum) the distal membranous part is unusually elongate and whip-like
(nearing twice as long or so as the sclerotized length, as in Text-fig. 38). The
epandrium (To) is very large and shows some very minor differences in shape between
species, but the differences are rather too intangible to have practical value in taxo-
nomy. The cerci are elongate, not fused, and sometimes show differences in shape
either in profile or posterior view which are useful for species recognition; they have ©
no features of use for supraspecific characters. The most important structures by —

REVISIONARY CLASSIFICATION OF RUTILIINI 17

far in the genitalia for taxonomic purposes are the surstylz ; these show great diversity
of shape, ranging from sharply pointed rather hook-like structures to enormous
foliaceous lobes. In some groups the surstylus shape may be very constant (e.g. in
Donovanius) throughout a range of species, but other groups may contain an admix-
ture of species some of which have surstyli differing only by subtleties of outline and
others of which have unmistakable surstyli showing some bizarre features. Broadly
speaking, however, each genus-group segregate tends to show a surstylus shape with
some common denominator—for example a surstylus always ending in a sharp tip or
always with a subquadrate form.

Female postabdomen. This has not been studied in detail, but the examination of
the postabdomen from representative females of different genera has not suggested
the existence of good characters of value in supraspecific taxonomy. Detailed
examination of species within a particular subgenus or genus will almost certainly,
however, show up some differences that will enable females to be more reliably
determined than at present.

KEY TO TRIBES OF THE SUBFAMILY PROSENINAE IN THE
ORIENTAL AND AUSTRALASIAN REGIONS

The subfamily Proseninae (=Dexiinae of authors) contains a vast assemblage of
Tachinidae that occur in all the zoogeographical regions and subregions (except
New Zealand) and appear to be exclusively parasites of beetles. If certain aberrant
fringe genera are excepted the Proseninae as a whole has a facies which—in spite of
great diversity in body shape and the degree of development of a facial carina—
specialists on Tachinidae can recognize, even if they find it hard to pin down exactly
what the diagnostic characters of the subfamily really are; certainly the form of the
maleaedeaigus andits associated structures seems to provide some common denomina-
tor throughout the group. At present there is no available recent definition of the
Proseninae, and the welter of world forms to be considered will make it difficult to
atrive at a satisfactory definition that will work on a world basis. Likewise the
completely satisfactory delimitation of tribal groupings within the subfamily will not
be easy, and Townsend’s various tribes in his Manual of Myiology need a thorough
revision. In these circumstances I am not attempting here to provide a definition
of the subfamily Proseninae, but I nevertheless think it useful to indicate the tribal
entities which seem to justify recognition in the fauna of the Oriento—Australasian
regions (in the area to which the Rutiliini are confined) ; an attempt will be made in a
later work on the Tachinidae of New Guinea to elaborate complete diagnoses of the
tribes and of the Proseninae as a whole.

The three tribes recognized in the Oriental and Australasian fauna are the
Prosenini, Doleschallini, and Rutiliini, which can be distinguished by the following
key.

1 Thorax closed above the hind coxae by a broad sclerotized bridge (as in Cylindro-
mylini), the hind coxae remote from the abdominal base. Head in profile sub-

triangular, profrons extraordinarily prominent and lower part of head strongly
receding, head very much longer at antennal axis than at epistomal axis. No

18 R. W. CROSSKEY

facial carina. Notopleuron not differentiated from prescutum by any depression.
Abdominal Tir + 2 excavate only at base. Body and legs excessively long and
slender, the long thin abdomen with subparallel sides. [Ceylon, Indonesia to
Solomon Islands, unknown from Australia] : : . Tribe DOLESCHALLINI
[Type-genus: Doleschalla Walker, 1861]
- Thorax membranous or mainly so between bases of hind coxae and abdominal
insertion, coxae and abdomen usually not remote from each other. Head in
profile not subtriangular, at most only slightly shorter at the epistomal axis than at
the antennal axis, often with epistome at least as prominent as profrons. Head
usually with facial carina (absent in some forms). Notopleuron distinctly differen-
tiated from prescutum by a groove or at least a shallow depression. Excavation
of abdominal Tr + 2 reaching to hind margin. Body form often robust, if long
and slender then abdomen broadest near base or subfusiform . 2
2 Postalar callus with only the normal two strong setae (at most only a shore weak
setula in addition). Suprasquamal ridge and postalar wall bare. Epistome not
produced in front of vibrissae or only weakly so, not readily visible in profile.
Propleuron bare or haired. Barette bare or with a few hairs anteriorly (if all haired
then no facial carina). Facial carina present or absent. Scutellum with three
pairs of marginal setae. Male hypopygium without long strong erect setae on
T7 + 8 except in Acuceva. Prosternal membrane and prosternum bare. Arista
often very long-plumose. [nearly cosmopolitan] . . Tribe PROSENINI
(Type-genus: Prosena Lepeletier & Serville, 1828]
— Postalar callus with supernumerary strong setae, total of 3-6 setae (except in
Chetogaster!). Either suprasquamal ridge or postalar wall usually haired.
Epistome subnasute or at least slightly and distinctly projecting in front of vibrissae
in profile. Propleuron haired (very rarely bare in isolated specimens). Barette
thickly haired along its length (partially bare in Chetogastey1). Strong facial
carina present. Scutellum with four or more pairs of marginal setae (except in
Chetogastey1). Male hypopygium with long strong erect setae on T7 + 8 (usually
standing in line on each side of tergite). Prosternal membrane, or membrane and
prosternum itself, sometimes haired. Arista micropubescent to short-plumose
(never very long plumose). [Oriental and Australasian Regions (except New
Zealand)] . 3 : : : : Tribe RUTILIINI
[Eype- genus: Rutilia Robineau-Desvoidy, 1830]

Tribe RUTILIINI Brauer & Bergenstamm

RUTILIAE Swainson im Swainson & Shuckard, 1840: 377. Generic plural unavailable as
family-group name (Article 11 (e) (i) of International Code of Zoological Nomenclature, 1961).

RUTILIIDAE Brauer & Bergenstamm, 1889: 76, 152. Type-genus: Rutilia Robineau-
Desvoidy, 1830.

AMPHIBOLIIDAE Brauer & Bergenstamm, 1889: 76, 152. Type-genus: Amphibolia Mac-
quart, 1843.

ROEDERIIDAE Brauer & Bergenstamm, 1889 : 76, 152. Type-genus: Roederia Brauer &
Bergenstamm, 1893 [=Chrysopasta Brauer & Bergenstamm, 1889], junior homonym of
Roederia Mik, 1881.

SENOSTOMATINI Townsend, 1932 : 40; Townsend, 1936 : 20, 154; Townsend, 1938 : 424.
Type-genus: Senostoma sensu Townsend, not Macquart [misidentification] [=Prodiaphania
Townsend]. SENOSTOMINI Enderlein, 1936 : 397, 435.

AGALMIINI Enderlein, 1936 : 397, 433. Type-genus: Agalmia Enderlein, 1936 (junior

1 This genus is rather intermediate between Rutiliini and Prosenini but is here retained in its tradi-
tional position in Rutiliini.

REVISIONARY CLASSIFICATION OF RUTILIINI 19

homonym of Agalmia Enderlein, 1934) [=Gvapholostylum Macquart, 1851, by subjective
synonymy of type-species].

HABROTINA Enderlein, 1936: 398 (subtribe). Type-genus: Habrota Enderlein, 1936
[=Chrysorutiia Townsend, 1915, by junior objective synonymy].

Diacnosis. Facial carina well developed, usually broadly separating antennae. Antennae
small, often falling short of mouth-margin by more than their own length; arista micropubescent
to short-plumose. Epistome projecting at least slightly, usually very strongly and sharply,
in front of vibrissae in profile, about as prominent as or more prominent than profrons. Vibris-
sae often reduced. Eyes bare, head in § sometimes nearly holoptic. ¢ frons narrower than 2
frons; proclinate orbital setae always absent in gj, weak or absent in 2. Proboscis short, not
longer than head; mentum in profile parallel-sided or slightly tapering. Scutal chaetotaxy
variable, often reduced. Scutellum, venter of sternopleura and mid and hind coxae often with
stiff spiniform setae. Prosternal membrane (in Chrysorutilia anterior angles of prosternum
also) sometimes haired. Propleuron haired (very rarely bare in isolated specimens). Barette
usually fully haired. Infrasquamal setulae absent (except in some Chetogastey species).
Postalar callus with 3-6 strong setae (except only two in Chetogaster). Postalar wall some-
times haired. Presutural intra-alar setae absent, 0o—3(4) post ia setae. Suprasquamal ridge
bare or haired. Lower calyptrae broad and abutting close to scutellum. Wing node with

small inconspicuous setulae above and below, wings otherwise bare. Second costal sector
bare below. Cell Rs open. Bend of vein M without appendix, very close to wing margin.
Vein Sc meeting costa just basad of or level with ym. Wings often with large dark area over
basal cells. Abdomen very large, broad and widest near base, posterolateral corners often very
prominent and T5 then often with median depression; T1 + 2 excavate to hind margin.
Abdominal chaetotaxy often spiniform. Tr + 2 without median marginal setae. T7 + 8 of
6 hypopygium with a group of long strong setae (usually 3 or 4 standing in line) on each side.
Medium-sized to very large flies (length 6-22 mm), often brightly coloured and metallic or with
conspicuous patterning.

IMMATURE STAGES AND BioLtocy. Parasitic in large soil-inhabiting white grub
larvae of Scarabaeidae (Melolonthinae, Rutelinae, Dynastinae) and in rotten-wood
inhabiting larvae of Lucanidae: probably also parasitizing other related Scarabaeoid
beetles. Eggs medium macrotype, slender, elongate and slightly bowed with
rounded ends (Townsend, 1942, plate 21, figs 146 & 147). Habit larviparous;
uterus enlarged, capacity of many hundred first stage larvae, these active and
deposited on soil-surface. Stage I larva slender, elongate, with long terminal hairs,
sometimes also some marginal segmental hairs; stage II larva unknown; stage III
larva swelling towards posterior end, segmentation distinct, cuticle (in one species
known) with covering of short colourless stubby hairs, posterior spiracles in form of
very large slightly separated plates either flush with posterior surface or slightly
sunken, plates perforated by very numerous minute pores or sinuate micro-slits,
outer rim of plates thickened, button subcentral. Puparium (of the few known
forms) with sunken posterior spiracles and covering of micro-hairs. Adult flies
absent from desert areas, present in scrub or forest, settling on tree-trunks, posts, or
underside of leaves in rain forest, females resting on ground only when larvipositing;
attracted to flowers, notably Eucalyptus.

DisTRIBUTION. Widespread throughout, and confined to, the Oriental and
Australasian zoogeographical regions (map, p. 163), but especially abundant in
Australia. In Oriental Region occurring through south-east Asia from Ceylon and
India to Philippines and Indonesia; in Australasian Region occurring in Australia

20 R. W. CROSSKEY

and Tasmania, Moluccas, Timor, Aru Islands, Kai Islands, New Guinea, Solomon
Islands, New Hebrides, Fiji, Samoa, and Lord Howe Island. Unrecorded from New
Caledonia, but probably occurring there. Absent from New Zealand.

Discussion. Since the time of Brauer & Bergenstamm (1889) specialists in the
Tachinidae have been agreed in regarding the Rutiliines as a named family-group
segregate distinct from other Proseninae (=Dexiinae of authors), though the group
has been variously ranked as a tribe, subfamily, or even occasionally as a family, and
sometimes has even embraced the Ameniines—a curious group of calyptrate flies
with an astonishing superficial likeness to the Rutiliines but now accepted as a sub-
family of Calliphoridae (Crosskey, 1965). It is unnecessary here to review all of the
varying interpretations of status and scope of the group, but the few major works of
the past 35 years require brief consideration.

Townsend (1936) treated the one family Tachinidae, as currently accepted, as
being seven families, of which the Rutiliidae were a small family on their own, placed
between the Prosenidae and the Tachinidae in Townsend’s narrow sense (Townsend,
1936 : 20, 150-156; 1938 : 410-427); Townsend’s families are, however, very un-
satisfactorily defined, and his family key has 84 exits (of which three are to Rutiliidae)

for the separation of seven families. One of the exits to Rutiliidae in the key ©
(Townsend, 1936 : 8) relates to those Rutiliines which have the suprasquamal ridge _

(=tympanic ridge of Townsend) haired, and there is never any problem in distin-
guishing these forms from all other Tachinidae: but it is much more difficult in
defining the Rutiliines to take account of the forms which, though obviously Ruti-
liines also, have the suprasquamal ridge bare—for the characters of the Rutiliines
as a whole then merge rather imperceptibly with those of the Prosenines (Prosenidae

of Townsend). For this reason it is impossible to separate the Rutiliines from the _
Prosenines so distantly as to rank the groups equally (as Townsend does), and it is —

much better therefore to treat the Rutiliines as being a subgroup within the Prose-
nines (Dexiines of authors).

This view was well expressed by Mesnil (1939), in his general essay on Tachinid
classification, as he divided the subfamily Dexiinae (Proseninae) into two tribes, —
Rutiliini and Dexiini (Prosenini), distinguishing the former from the latter by the ©
prominent mouth-margin together with a few other less tangible features. My own —

view is very much in accord with that of Mesnil (1939), both as regards the affinities
and ranking of the Rutiliines, and is elaborated in more detail in the following
discussion (in which I have used the currently accepted names Proseninae and
Prosenini in place of Dexiinae and Dexiini as used by many authors).

The vast complex of diverse forms which constitute the subfamily Proseninae is

exceedingly hard to classify satisfactorily at any level, and almost any constellation
of characters which appears to be satisfactory for defining a genus-group or family-_

group taxon when only a small collection or a limited regional fauna is available
tends to become almost worthless for taxon definition when a comprehensive collec-
tion of worldwide forms is studied. Yet so great is the number of forms to be coped
with that some segregation of family-group categories is needed, even if these are

difficult to define in a fully satisfactory way and even if some genera are hard to

REVISIONARY CLASSIFICATION OF RUTILIINI 2

place into a subtribe or tribe with any confidence. Considering the whole of the
Proseninae there is no doubt that the great bulk of forms constitute one very large
tribe (Prosenini) containing much variety of form but united by all perceptible shades
of intermediate characters, but that there are some other large groups or small
groups that are more disjunct from the main body of forms and which can justifiably
be ranked as tribes equivalent to the Prosenini; these include the Rutiliini, Doleschal-
lini and Trixini, and perhaps a few other definable tribes when sufficiently studied.
The Theresiini, Dexillini and Zeliini which Townsend (Manual of Myiology) treated
as distinct tribes appear to me to be indistinguishable from Prosenini, and I am also
doubtful whether the Trichodurini can be separated from the Prosenini either.
The position of the subtribe Stominina of Mesnil (1939 : 52-53) is also very doubtful,
but in my view it has scarcely any of the characters of the Rutiliini and cannot be
placed in the Rutiliines where Mesnil has classified it: the characters of Stomina
Robineau-Desvoidy appear to place this genus close to Billaea Robineau-Desvoidy
in the Prosenini. At present the various tribal segregates within the Proseninae are
insufficiently clarified, but the diagnosis here given should certainly distinguish the
Rutiliini satisfactorily from all other Proseninae (except perhaps for the anomalous
genus Chetogaster), and the key given in a foregoing section will separate the tribe
from other tribes of Proseninae found in the Oriento—Australasian regions.
Although the Rutiliines have been so widely accepted as a family-group segregate,
and are here ranked as a tribe on their own, it is impossible to characterize them
as a whole by any simple character or small group of characters in a way that
absolutely defines them. The best diagnostic feature (though it does not hold for
| the genus Chetogaster) is the presence of supernumerary strong bristles on the postalar
callus ; in all other Proseninae (and virtually all Tachinidae) there are only two strong
setae on the postalar callus, sometimes accompanied by a long hair, but ia all
Rutiliini (excepting Chetogaster) there are at least three very strong postalar setae
and sometimes from four to six. Another important character is the snout-like
| development of the epistome, which projects far in front of the vibrissae in profile
| (as mentioned by Mesnil, 1939), but some Formosia and Rutilodexia have the epistome
| only very slightly projecting, and resemble many Prosenines in this respect, so that
this character, too, is not completely satisfactory. Many Rutiliini have the supra-
squamal ridge thickly haired and others have a tuft of long hair standing on the
wall of the postalar callus, and these features are never found in other Proseninae.
The facial carina is always very strong in Rutiliini, and often very wide and flattened
(then assuming a form almost never found among other Prosenines), and many
members of the tribe have very strong erect spiniform setae on the abdomen or long
| strong spiniform setae on the scutellum and mid and hind coxae; such spiniform
setae are of rare occurrence in other Proseninae, although they are found in several
South American genera and in the Nearctic genus Euchaetogyne Townsend, but the
New World forms which slightly resemble Australasian Rutiliini can always be
easily distinguished by possessing only the two normal setae on the postalar callus
and usually by having the propleuron bare (though in Euchaetogyne the propleuron
is haired, asin the Rutiliini). The males of all Rutiliini have some very characteristic
long strong bristles on tergite 7 + 8 of the hypopygium, these bristles standing in

22 R. W. CROSSKEY

two groups separated medially and normally consisting of three or four such setae
standing in a transverse line on each side; long strong setae of this kind on T7 + 8
almost never occur in other Proseninae, though similar setae are found in the Euro-
pean genus Deximorpha Rondani and the Australian genus Acucera Malloch.

Enderlein (1936) divided the Rutiliines, which in his treatment ranked as a sub-
family, into five tribes (of which two tribes are Ameniines), and Townsend (1936,
1938) divided his Rutiliidae into two tribes, but both of these workers are renowned
for their taxonomic ‘splitting’ and later workers have, rightly in my view, declined
to accept any validity for these tribes; nor have the many ill-defined and unnecessary
genera erected by Enderlein (1936) been accepted (Paramonov, 1968 : 351, has
pointed out that they are unwarranted, and often misleading because of misidentifi-
cation of the type-species, and I agree entirely with Paramonov’s view). In the
present work no subtribes are recognized, and the scope of the Rutiliini is very
similar to that shown by Paramonov (1968), except that the genera Ola Paramonov,
1968, and Ruwya Paramonov, 1968 (of which the type-species and several other
included species have been examined) are excluded: the facies of Ola and Ruya are
not at all those of Rutiliini, both possess three strong post ia setae (such as rarely
occur in Rutiliines), the conformation of the flat face is quite unlike Rutiliines, and
there are many other features on which Ola and Ruwya must be excluded from the
tribe. It should be noted that Paramonov (1968 : 381) was himself doubtful about
the inclusion of these genera in Rutiliini.

Paramonov (1968 : 350-352) has given an historical review of the description of
Rutiliine genera, which need not be repeated here. Up to now 38 generic and
subgeneric names have been proposed for Rutiliine flies, most of which are nomen-
claturally available though many of them are considered taxonomically invalid in
Paramonov’s (1968) treatment and in the present work. Two names published by
Paramonov (1968), viz. Formotilia and Rutilosia, are unavailable under Article 13 (b)
of the International Code of Zoological Nomenclature because there is no fixation of a
type-species for either genus. The names Diaphania Macquart, Roederia Brauer &
Bergenstamm, Agalmia Enderlein and Ewucompsa Enderlein are preoccupied
homonyms, and the replacement name Prodiaphania Townsend is available for the
first of these; the others are synonyms and do not require replacement names.
The name Eucompsa Enderlein, 1936, in Rutiliini was proposed by Enderlein (1936 :
400) as a new genus, with Rutilia minor Macquart cited as type-species, although
Enderlein had himself already erected the genus Eucompsa Enderlein, 1922, in
Tabanidae, and R. minor was already type-species of Microrutilia Townsend: thus
in one name Enderlein achieved the feat of publishing a junior objective synonym
(of a Townsend genus) and a junior homonym (the latter of one of his own names)!

In the present re-classification of the Rutiliini I recognize the following eight
genera: Formosia Guérin-Méneville, Rutilodexia Townsend, Formodexia gen.n.,
Rutilia Robineau-Desvoidy, Amphibolia Macquart, Chrysopasta Brauer & Bergen-
stamm, Prodiaphania Townsend, and Chetogaster Macquart. Three of these are
divided into subgenera, three subgenera being recognized in Formosia, seven in
Rutilia and two in Amphibolia.

A detailed study of the male genitalia throughout the tribe has shown that there

Al,

REVISIONARY CLASSIFICATION OF RUTILIINI 23

are no characters that can be used at the generic level (although differences in the
shape of the surstyli and cerci can be very important in distinguishing species) and
genera have to be characterized by externally visible features. Some of the genera
are moderately distinctive and can be satisfactorily diagnosed, but others need to be
characterized by aggregates of several features taken together (even then they may
have to include species which clearly fit them on total facies but which fail to conform
to one or more of the normal generic characters). It has to be borne in mind when
using the keys that almost any character can fail in an occasional specimen.

KEY TO THE GENERA OF RUTILIINI

[Note : Tasmania is treated as part of Australia and is not separately cited in the geographical
information. ]

I Suprasquamal ridge bare (Text-fig. 21). Prosternal membrane bare : 2
— Suprasquamal ridge haired (Text-figs 23 & 24). Prosternal membrane =e or
haired (Text-fig. 18) : 7

2 Palpi minute, not longer than basal width of mentum, and not longer than third
antennal segment. Buccal opening very long and narrow (Text-fig. 15), at
narrowest point not wider than facial carina. Arista conspicuously short-plumose.

Upper parts of parafacials haired. Upper calypter enlarged, when wings folded
usually as long as lower calypter. [Australia only]
PRODIAPHANIA Townsend (p. 107)

— Palpi well developed, very much longer than either basal width of mentum or third
antennal segment. Buccal opening wide (Text-fig. 14), much broader than facial
carina (except in Forvmodexia). Arista pubescent or occasionally with very short
plumosity. Parafacials entirely bare (some exceptions). Upper calypter normal 3

3 Postalar callus with two strong setae (sometimes with one very much shorter and
finer setula in addition differentiated from hair). Epistome subnasute, facial
profile deeply excavate between epistome and carina (Text-fig. 8). Palpi sexually
dimorphic, distinctly clubbed or spatulate in 2 and slender in g. Infrasquamal
setulae often present. Scutellum normally with only three pairs of marginal setae,
at most with four (including the apical pair). [Australia only]

CHETOGASTER Macquart (p. 112)

— Postalar callus with three or more strong setae. Epistome not very strongly promi-
nent, facial profile usually only slightly concave between epistome and carina
(Text-fig. 4). Palpi not sexually dimorphic, long and slender in both sexes (at
most only trace of swelling at extreme tip). Infrasquamal setulae absent.
Scutellum normally with 4-11 pairs of a setae (including apical pair), very
rare specimens with only three pairs : 4

4 Apical scutellar setae inserted at the same level as the other ecutellar neampinal sotae
(Text-fig. 11). Postalar wall with dense hair tuft (Text-fig. 25). Ventral margins
of abdominal tergites with very strong spiniform setae directed downwards. Head
often partly metallic. g head sometimes holoptic or almost so. [Java to Solomon
Islands, Australia] ‘ : . FORMOSIA Guérin-Méneville (p. 25)

— Apical scutellar setae inserted at a conspicuously lower level than the other scutellar
marginal setae (Text-fig. 10). Postalar wall without dense hair tuft (except in
Formodexia). Ventral margins of abdominal tergites without such vestiture, if
marginal setae slightly spiniform then directed backwards as well as downwards.

Head non-metallic. g head never holoptic ; 5

5 Postalar wall with dense hair tuft. Buccal opening very narrow, in rg scarcely at all

24 R. W. CROSSKEY

wider than facial carina. Tegula without long fine wirysetulae. Palpi exceptional-
ly long and slender. Costal base explanate and with very long dense fringe
(wings in folded position therefore appearing to have strongly developed
‘shoulders’). [Moluccas only]. : . FORMODEXIA gen. n. (p. 38)
- Postalar wall without hair tuft, money, eaeey bare (few specimens with a few hairs
on extreme upper part adjacent to the dorsolateral ridge of the postalar callus).
Buccal opening broad, very much wider than facial carina in both sexes. Tegula
with normal long fine wiry posterior setulae. Palpi normal. Costal base not
explanate (wings in folded position without appearance of prominent ‘shoulders’).
[Not from Moluccas] . 6
6 Genal dilation small, only extending forwards for Boone fale the iene = ihe genal
region and not as far as the most anterior point of the eye (Text-fig. 5). Proboscis
with slender mentum that is conspicuously tapering in profile (Text-fig. 13).
Postalar callus with 4-6 setae and body without white pollen spots on sides of
thoracic dorsum and the abdomen. Wing membrane distinctly tinged with
brown or yellowish brown. [New Guinea, New Britain, Aru Islands]
RUTILODEXIA Townsend (p. 40)
— Genal dilation well developed, reaching forwards for obviously more than half the
length of the genal region and as far as the most anterior point of the eye (Text-
fig. 4). Proboscis with broader mentum in which sides subparallel in profile
(Text-fig. 12). Postalar callus with either three strong setae only or with 4—5 setae
and body with white pollinose spots on thorax and abdomen. Wing membrane
(except for basal dark mark) clear hyaline or almost so. [Australia]
RUTILIA Robineau-Desvoidy (a few forms)
7 Dorsum of thorax with four broad bold black vittae which are interrupted at the
transverse suture, appearing therefore to have eight elongate black spots (four on
prescutum and four on scutum). Palpi much shorter than mentum. Postorbits
with alternating silvery white and black spots which shift in appearance with
direction of light. Facial carina strongly bulbous on upper part and strongly
contracted with sharp median ridge on lower part. Parafacial hairing reaching
below the lowest point of the eye and virtually continuous with hairing on genal
dilation. [Western Australia only]
CHRYSOPASTA Brauer & Bergenstamm (p. 102)
— Dorsum of thorax without such pattern. Palpi as long or almost as long as mentum.
Postorbits without pattern of alternating white and black areas. Facial carina
broad and flattened on anterior surface with subparallel sides, or widened above and
contracted ventrally (in latter case not normally formed into very narrow sharp
ridge). Parafacials bare or haired, but if haired the hairing not normally extending
below lowest point of eye in profile and well separated by conspicuous bare area from
hairing on the genal dilations : : : : 8
8 Intermediate abdominal tergites (T3 and T4) pace Giscal seeae! Facial carina
usually very broad and flattened, often slightly sulcate, with subparallel sides, at
most only a little knob-like dorsally and contracted ventrally. Abdomen without
bold black-and-white pattern (rare exceptions). Forms often with partly metallic
head or with deep median depressioninabdominalT5. [Widespread in Oriental and
Australasian Regions] . : RUTILIA Robineau-Desvoidy (most forms) (p. 42)
— Intermediate abdominal tergites (T3 and T4) with discal setae (lacking in occasional
specimens on one or other of these tergites, very rarely on both; if absent then
abdomen with bold black-and-white pattern). Facial carina widest dorsally
between junctions of second and third segments of the antennae and conspicuously
narrowed ventrally, not subparallel-sided and not flattened on anterior face.
Abdomen most often with bold black-and-white pattern. Head never metallic,
completely pollinose, and abdominal T5 without wide median depression. [New
Guinea, Australia and Lord Howe Island] ; . AMPHIBOLIA Macquart (p. 93)

REVISIONARY CLASSIFICATION OF RUTILIINI 25

Genus FORMOSIA Guérin-Méneville

Formosia Guérin-Méneville, 1843 : 263. Type-species: Rutilia mirabilis Guérin-Méneville
1831, by monotypy.

Dracnosis. Facial carina large, widening ventrally or with subparallel sides, at most only
slightly widened medially. Epistome usually not strongly prominent. ¢ head holoptic or
nearly so, eyes always strongly approximated, upper eye facets sometimes conspicuously en-
larged and set off from small lower facets. Parafacials always bare. Buccal opening normal,
if somewhat narrowed then conspicuously wider than facial carina. Genal dilation well
developed, usually not reaching forward as far as front level of eye. Head often partly metallic.
Arista pubescent or short-plumose. Palpi normal. Prosternum and prosternal membrane
bare. Scutellum with apical setae at same level as, and not readily distinguishable from, other
marginal setae; marginal setae stiff and straight, normally at least five pairs; disc of scutellum
often markedly flattened. Postalar callus with 3-5 strong setae. Postalar wall with dense
hair tuft. Suprasquamal ridge bare. Upper calypter normal. Tegula with normal long wiry
posterior setulae. Costal base not explanate. Abdomen with stout spiniform setae directed
downwards on ventral margins of tergites; T3 with median transverse row of spiniform marginal
setae (except in males of most Euamphibolia spp.); intermediate tergites without discal setae.
T5 short and broad, usually with well formed median depression and prominent posterior
corners.

DISTRIBUTION. From Java to Solomon Islands and Queensland, one species (see
Euamphibolia) reaching to Tasmania and Western Australia; especially well repre-
sented in Moluccas, New Guinea and Bismarck Archipelago. Unknown from
Philippine Islands and possibly absent.

Discussion. This is the predominant genus of Rutiliini in the Papuan subregion
and contains many large and attractive species of beautifully patterned and metallic
flies which often have a brilliant golden green to violet-blue coloration; the general
appearance, both of shape and colour patterns, resembles that of many Rutilia
species, and Formosia and Rutilia are without doubt very closely allied genera.
Despite the superficial resemblance, Formosia is easily distinguished from Rutilia,
and from all other genera of Rutiliini here recognized (except Formodexia gen.n.) by
the presence of a thick tuft of long hair on the side wall of the postalar callus (Text-
fig. 25). This is a remarkable and very unusual character which, to the best of my
knowledge, does not occur elsewhere in the Tachinidae, and in the Rutiliini is
confined to Formosia and Formodexia (though occasional specimens of Rutilodexia
and Rutilia s.1. may have a very few hairs on the extreme upper edge of the postalar
wall adjacent to the postalar rim). The features distinguishing Formosia from
Formodexia are discussed under the latter genus, and a tabulation of the main
differences between Formosia and Rutilia s.l. is given in the treatment of Rutilia.

Six genera have been described in the Rutiliini which are here considered not to be
generically distinguishable from Formosia. These are Pseudoformosia Brauer &
Bergenstamm, Euamphibolia Townsend, Chromocharis Enderlein, Hega Enderlein,
Laccura Enderlein and Pancala Enderlein. Two of these names, however, apply to
rather well defined segregates within Formosia s.l. which it is thought useful to recog-
nize as subgenera, as these subgenera (viz. Pseudoformosia and Euamphibolia) can
be defined by quite constant structural features which are associated with rather
conspicuous differences in general facies and pattern. The other four names (all

26 R.-W. CROSSKEY

provided by Enderlein in his usual lavish way) are treated as unnecessary synonyms
applying to concepts for which older genus-group names are available: Chromocharis
and Hega are placed as synonyms of Euamphibolia, Laccura as a synonym of Pseudo-
formosia, and Pancala as a synonym of Formosia s.str.

The male genitalia and fifth abdominal sternite in Formosza s.l. call for no special
comment as they differin no significant way from those of other Rutiliini or Proseninae
in general. The shape of the surstyli (paralobes) is more square and ‘heavy’ in the
subgenus Evamphibolia than in the other subgenera of Formosia, and males of this
subgenus are therefore rather distinctive.

KEY TO THE SUBGENERA OF FORMOSIA

1 Postalar callus with 4-5 strong setae. One post ia seta (rarely none). Eyes of ¢
very strongly approximated or meeting, upper part of frons narrower than facial
carina. Upper eye facets of $ often greatly enlarged. Head partly metallic (a
few exceptions). No posthumeral setae. Mentum in profile normally rather
slender and distinctly tapering (Text-fig. 13). Surstyli of f genitalia of varied form
but normally tapering on apical half and not subquadrate (Text-figs 41-44) ‘ 2
- Postalar callus with three strong setae. 2-3 post ia setae (sometimes very weak
fourth seta). Eyes of $ not very strongly approximated, never meeting, frons at
narrowest at least as wide as and normally wider than the facial carina. Upper eye
facets of g not enlarged. Head non-metallic, entirely pollinose. Small post-
humeral seta normally distinguishable among the prescutal hairing. Mentum in
profile not unusually slender, upper and lower edges subparallel (Text-fig. 12).
Surstyli of 3 genitalia very heavy and subquadrate in profile (Text-figs 52 & 53)
EUAMPHIBOLIA Townsend (p. 33)
2 Anterior surface of fore coxa almost completely haired (Text-fig. 16). One sterno-
pleural seta (o + 1). Presutural dorsocentral setae absent. Abdominal T5 with-
out long strong setae behind the main transverse row
FORMOSIA Guérin-Méneville s.str. (p. 26)
- Anterior surface of fore coxa bare on inner half, except near apex (Text-fig. 17).
Two sternopleural setae (1 + 1). At least one small presutural dorsocentral seta
present on each side of prescutum, though sometimes very weak. Abdominal T5
with or without long strong setae behind the main transverse row
PSEUDOFORMOSIA Brauer & Bergenstamm (p. 31)

Subgenus FORMOSIA Guérin-Méneville

Formosia Guérin-Méneville, 1843 : 263. Type-species: Rutilia mirabilis Guérin-Méneville,
1831, by monotypy.

Pancala Enderlein, 1936: 422. Type-species: Formosia callipygos Gerstaecker, 1860, by
original designation. Syn. n.

Diacnosis. Head metallic, at least in some lights, on postorbits, genal dilation, parafrontals
and usually also upper parafacials (except in vividiventris sp.n.). Head of g holoptic or nearly
so, upper interfrontal area obliterated or nearly so by meeting parafrontals, frons at its narrowest
point not as wide as facial carina. Upper facets of g eyes often much enlarged, ocelli then
very prominent. Arista shortly plumose, hairs longer than basal thickness of arista. Mentum
of proboscis rather slender, tapering distally. No posthumeral setae. No presutural dorso-
central setae. One strong post ia seta. One sternopleural seta (o + 1). Postalar callus with
4-6 setae. Scutum often with supernumerary strong prescutellar setae between the hindmost
post acy and post dc setae, the setae together forming a transverse prescutellar row. Disc of

REVISIONARY CLASSIFICATION OF RUTILIINI 27

scutellum usually conspicuously flattened. Fore coxa almost completely haired on anterior
surface. Transverse row of erect median marginal setae of T3 present in both sexes. T5 with-
out strong setae behind the transverse row (a few exceptions).

DISTRIBUTION. Occurring from Malaya and Java eastwards to the Solomon
Islands, and known from Selangor, Java, Celebes, Molucca Islands (including Halma-
hera, Ternate, Batjan, Morotai, Obi, Seram), New Guinea, Aru Islands, New Britain,
New Ireland, Lavongai, Bougainville, and the major islands of the Solomons chain
south-east to Guadalcanal. Unknown from Philippine Islands (possibly not
represented there), absent from Australia.

Discussion. Formosia s.str. is the largest subgenus in number of described
species (and several additional undescribed species are known from the New Guinea
area) and contains most of the brilliant metallic forms of Rutiliini that form such a
characteristic element in the Tachinid fauna of the Papuan subregion. The sub-
genus is easily distinguished by the extensively haired fore coxae, in which almost
all of the anterior surface is haired (Text-fig. 16), an unusual feature found nowhere
else in the Rutiliini; in other Rutiliines, as in most Tachinidae, the inner half of the
anterior surface of each fore coxa is almost entirely bare (there being only a few hairs
towards the tip). The one sternopleural seta and the one posterior intra-alar seta,
together in correlation, also uniquely distinguish Formosia s.str. from the other
subgenera. A rather unusual character, too, though occurring in some other Ruti-
liini, is the slender tapering proboscis, and the strongly flattened scutellum of most
forms is another feature exceptional among Rutiliini as a whole.

Formosia s.str. is especially well represented in the lowland rain forests of New
Guinea and the Bismarck islands, where some species are common at times, but a
few species also occur in small patches of relict forest in the New Guinea highlands.
Adult flies can often be found resting on low herbage, especially on the undersides
of the leaves of wild ginger and similar plants, but nothing is yet known of the host
relations (almost certainly, however, Scarabaeoid beetles are the hosts).

Nearly all the species of Formosia s.str. form a rather homogeneous group having
characters closely similar to those of the type-species, PF’. (F.) mirabilis, but a new
species (described herein) from Guadalcanal, in the Solomon Islands, is rather dis-
junct from the main body of species—although it fits Formosia s.str. well in all
essential characters—and is here treated as a group of its own. The two groups
recognized are keyed and defined below.

KEY TO THE SPECIES-GROUPS OF FORMOSIA s:.str.

1 Abdominal T5 with some long strong erect setae laterally behind the transverse row.
Head entirely thickly pollinose, nowhere metallic. Surstyli of ¢ genitalia in
profile strongly acuminate and ending in sharp point (Text-fig. 49), the whole
surstylus somewhat hook-like : . viridiventris-group (p. 28)
- Abdominal T5 without strong setae chine the transverse row of setae. Postorbits
and genal dilations, sometimes also parafrontals and upper parafacials, metallic.
Surstylus of § genitalia not of this form, always rounded at tip (Text-figs 41-44)
mirabilis-group (p. 29)

28 R. W. CROSSKEY

Tue VIRIDIVENTRIS-GRovupP

Diacnosis. Facial carina unusually narrow, in facial view subequal in width to an antenna.
Head entirely pollinose, no bare metallic areas. Thorax and abdomen unicolorous dark green,
no pattern. Abdominal T5 with some long strong setae laterally behind the transverse row
of setae. Surstylus of ¢ genitalia sharply pointed (Text-fig. 49).

INCLUDED SPECIES

Formosia (Formosia) viridiventris Crosskey sp. nm. GUADALCANAL. [Des-
cribed below].

Formosia (Formosia) viridiventris sp. n.
(Text-fig. 49)

6. Head. Ground colour blackish, except upper part of facial carina and lunula more
reddish brown, interfrontal area velvety black-brown; head rather thickly yellowish white
pollinose, nowhere metallic but trace of greenish colour showing through pollinosity along
lines of postocular setulae. Parafrontal and genal hair blackish, occipital hair yellowish white.
Eyes strongly approximated but head not holoptic, upper facets not enlarged, upper part of
interfrontal area almost eliminated by coming together of parafrontals in mid line; frons at
narrowest point 0-06—-0:075 of head width, subequal to width of facial carina. Ocelli slightly
raised, ocellar setae weak hair-like. Facial carina long, very slightly fusiform, longer than the
epistome, facial profile only very slightly and evenly concave at junction of epistome and carina.
Gena broad, 0-33—0-38 of eye-height. Parafacial about twice as wide as third antennal segment.
Antennae dark reddish brown to blackish brown, more orange at junction of second and third
segments, falling short of mouth-margin by about length of third segment; third segment
2°6-3-0 times as long as second segment; arista very short plumose. Palpi dark brown or dark
reddish brown with slightly paler tips. Mentum tapering, rather slender. Thovav. Uni-
colorous very dark green or bluish green metallic, sometimes with violaceous tinges. Prescutum
in some lights showing trace of four narrow blackish vittae and thin covering of whitish
pollinosity, but these so inconspicuous that prescutum appears concolorous metallic greenish
with the scutum. Thoracic hair all black except for postalar tuft pale yellow and the sparse
hairing of the propleura yellowish white. Chaetotaxy normal, setae not markedly spiniform.
Scutellum not flattened; with four pairs of strong marginal setae, these usually immediately
preceded by some moderately strong horizontal preapical setae. Wings. Basicosta dark
yellowish brown or reddish brown darkened to blackish brown anteriorly. Wing membrane
pale smoky brownish with darker brown infuscation along the veins; infuscation conspicuous to
naked eye and appearing more yellowish brown basally. Calyptrae dark brown with brown
marginal hair. Legs. Black with entirely black hair; rather elongate and slender with femora
attenuate on apical halves. Fore coxa fully haired anteriorly; mid femur without or with one a
submedian seta; hind tibia with one strong ad seta, without ad fringe and without pd setae.
Claws very long. Abdomen. Dorsum concolorous with thorax, uniformly metallic dark green
or dark bluish green, venter violaceous towards the sides. Abdominal hairing entirely black;
hair short, fine, and recumbent. Six setae in the transverse marginal row of T3. T5 with some
long strong setae standing among the hair on each side behind the setae of the main transverse
row. Depression of T5 deep and conspicuous. T7 + 8 of hypopygium with only one or two
long fine setae on each side, and these not very clearly differentiated from the hair vestiture.
Genitalia (Text-fig. 49) with apical part of surstylus very strongly contracted, curved forwards
and sharply pointed at tip (somewhat hook-like when seen in profile). Measurements. Body
length 16-7-18-3 mm (mean 17:5 mm), wing length 14-7-16-5 mm (mean 15-6 mm) [4 specimens,
holotype with minimum lengths indicated].

2. Extremely similar to g except for usual sexual differences. Interfrontal area very dark
reddish brown, narrowing dorsally but width at minimum about twice as great as a parafrontal;

REVISIONARY CLASSIFICATION OF RUTILIINI 29

one pair of strong proclinate orbital setae. Vertex o-16 of head width. Antennae more
reddish than in g (in the specimen seen). Body length 15-7 mm, wing length 13-4 mm [one
specimen only seen].

MATERIAL EXAMINED
Holotype 3, SoLomon IsLANpDs: Guadalcanal, Tapenanje, 10-30 [printed as
10—32].1x.1953 (J. D. Bradley). In British Museum (Natural History), London.

Paratypes. Soromon IsLtanps: 2 g, Guadalcanal, Gold Ridge, 1-2000 ft,
21.ix.1958 (P. G. Fenemore); 1 3, Guadalcanal, Honiara District, 500 ft, 2.1.1955
(E. S. Brown); 1 9, Guadalcanal, Honiara District, Mt Austen, 13.vi.1954 (E. S.
Brown). All paratypes in British Museum (Natural History).

DISTRIBUTION. Known only from the island of Guadalcanal in the southern
Solomon Islands.

ArrFinitiges. A rather isolated and distinctive species (here placed in the separate
species-group defined above) identifiable at a glance by the uniformly dark green
body and infuscate wings. The frons and vertex in the female are unusually narrow
for the genus. No particular affinity with any other species is evident.

THE MIRABILIS-Grovup

Diaenosis. Facial carina broad, in facial view conspicuously wider than an antenna. Head
partly metallic, at least on postorbits and genal dilations, often also on parafrontals and upper
parts of parafacials. Thorax and abdomen not unicolorous, abdomen with conspicuous pattern
or if not then thorax or scutellum not concolorous with abdomen. Abdominal T5 without
strong setae behind the transverse row of setae. Surstylus of § genitalia of varied form (Text-
| figs 41-44) but not sharply pointed at apex.

_ INCLUDED SPECIES

Formosia (Formosia) blattina (Enderlein) comb. n. CELEBES. [Holotype
examined].
F. (F.) bracteata (Enderlein) comb. n. NEw IRELAND. [Holotype examined].
F. (F.) callipygos Gerstaecker. NEw GuinEA. [Holotype examined].
F. (F.) eos (Enderlein) comb. n. CELEBES. [Lectotype examined].
F. (F.) fervens (Walker). Motuccas. [Holotype examined].
F. (F.) flavipennis (Macquart). Java, Mataya. [Holotype examined].
F. (F.) gemmata (Enderlein) comb. n. NEw Britain. [Lectotype examined].
viridescens (Enderlein). [Lectotype examined].
F. (F.) glorificans (Walker) comb. n. NEw GUINEA. [Holotype examined].
pectoralis (Walker) syn. n. [Holotype examined].
fulvipes (Enderlein). [Holotype examined].
F. (F.) heinrichiana (Enderlein) comb. n. CELEBES. [Holotype examined].
F. (F.) heinrothi (Enderlein) comb. n. New Britain. [Holotype examined].
F, (F.) mirabilis (Guérin-Méneville). NEw Guinea. [Holotype examined].
plumicornis (Macquart).
F. (F.) solomonicola Baranov stat. n. SoLromon IsLanps. [Lectotype
examined].

30 R. W. CROSSKEY

KEry TO SPECIES OF THE SUBGENUS FORMOSIA s. str.

[Note: F. (F.) viridiventris is omitted as it is easily recognized from the separate species-group
characteristics and description already given. Specific limits are very uncertain at present
for the other species, i.e. those of the mirabilis-group, and it is not yet clear to what extent the
small differences in male genitalia provide dependable specific criteria. The following key is
therefore very tentative and is merely a preliminary one which runs out the supposed species
on the external characteristics shown by the types. Specimens exist in museum collections of
undetermined species, possibly new, which will not necessarily run out in the key. ]

1 Legs predominantly reddish yellow (at most only the tarsi blackish and some
reddish darkening apically on the femora). Pleural regions of thorax with
mainly pale yellow to golden orange hair. Epistome reddish yellow to pale
tawny reddish. Third antennal in orange (sometimes suffused with light
brownish apically) ‘ 2

— Legs black-brown to black (at most only apices of femora pale and tibiae sometimes
a little reddish). Pleural regions of thorax with black or yellow hair. Epistome
blackish brown to black. Third antennal segment dark reddish brownto black . 4

2 Eyes of g very strongly approximated, interfrontal area obliterated on the upper
frons, frons at narrowest point not more than 0-04 of head-width; upper eye
facets conspicuously enlarged. Mair of tibiae all black. Upper parts of para-
facials conspicuously metallic in almost any light . ; 3

— Eyes of f not very strongly approximated (separated by distance chou fare nes
as great as diameter of anterior ocellus), interfrontal area reaching to ocellar
triangle, frons at narrowest point o-o8—o-10 of head-width. Hair of tibiae
partly or largely golden red. Upper parts of parafacials rather thickly pollinose,
hardly appearing metallic from any viewpoint. [Java] . : ‘ F. flavipennis

3 Hair of fore coxae mainly golden orange (only the strongest vestiture black).
Extreme costal base with golden red hairing ventrally. Tarsi reddish yellow to
dark red, almost concolorous with remainder of legs. Crinkled hair of mesopleural
hind margin almost all golden orange. Frons of § at narrowest point subequal
in width to anterior ocellus, about 0-04 of head-width. [Celebes] . : F. eos

— Hair of fore coxae mainly black (some golden hair at base). Hairing of costal base
all black. Tarsi blackish, conspicuously darker than remainder of legs.
Crinkled hair of mesopleural hind margin almost all black. Frons of g§ extremely
narrow, eyes nearly meeting, frons at narrowest point not as wide as anterior

ocellus, only about 0-02 of head-width. [New ey ee : ; : F. glorificans
4 Hair of pleural regions of thorax all black : : : : : 5
— Hair of pleural regions of thorax pong pale yellow or golden ‘ : : : II
5 Forms from Celebes . - : : : 6
— Forms from Moluccas, New Guinea or Bismarck eRe
6 Calyptrae smoky brown with brown fringes. Postbuccal and genal hair ‘black.

Abdomen mainly a beautiful, brilliant, metallic blue; blue colour occupying most of
each side of tergites 3-5 but divided medially by a broad bold black median vitta;
hind margins of T3—T5 only narrowly black. Wings rather uniformly greyish
brown infuscate. ¢ eyes separated by a distance only about equal to twice width
of anterior ocellus, but interfrontal area not obliterated and reaching to anterior
ocellus (interfrontal area conspicuous therefore even at narrowest point of frons
and slightly wider at this level than a parafrontal) . . ‘ . F. heinrichiana
[This species is known only from the ¢ holotype]

— Calyptrae pale orange-yellow with yellow fringes. Postbuccal and genal hair pale
yellow. Abdomen without such blue-and-black pattern, tergites 3-5 each with a
purplish red transverse band that is not, or only very faintly, interrupted

REVISIONARY CLASSIFICATION OF RUTILIINI 31

medially; the purplish red colour not strongly papers with the broad black
tergite hind margins. [¢ unknown] - : : . F. blattina
[This species is known only from the 2 holotype)”

7 Abdominal pattern consisting of golden green, coppery green or reddish copper

metallic bands on T3—-T5 which strongly contrast with a black centre line and

black tergite hind margins; the metallic bands on each side usually nil has

arcuate, especially on T3 and T4. [Moluccas and New Guinea] - 8
— Abdomen rather uniformly dark, the inconspicuous pattern consisting of a wang

dark blackish green or slaty bluish band across each tergite from T3—T5 which is

not medially interrupted and only very feebly contrasts with the black hind

margin of the tergite; bands not at all arcuate. [Bismarck Archipelago] . : 10
8 Calyptrae pale yellowish brown with pale yellow fringes; long fringe hair of outer

side of upper calypter conspicuously yellow. Wing bases yellow-brown to naked

eye. [Moluccas] F g . F. fervens
— Calyptrae brown or dark inca with pale own to dark brown fringes; long outer

fringe hair of upper calypter quite distinctly brown. Wing bases dark brownish

to naked eye. [New Guinea] : 9
9 Metallic fascia on each side of T5 with its hind es excarate and therefore nar-

rower on the centre part than at the inner end (the metallic band only reaching

back to the level of the transverse row of setae at the inner end) : F. mirabilis
— Metallic fascia on each side of T5 not noticeably excavate on the hind margin,

extending back as far as, or almost as far as, the transverse row of setae (metallic

band not wider at its inner end than elsewhere) : : F. callipygos
to Mesonotum and scutellum golden green to blue-green. Basicosta yellowish on
posterior part and only infuscate anteriorly. [New Britain] . : F. gemmata

— Mesonotum and scutellum mainly metallic coppery red or coppery purple, some
coppery green colour on edges of prescutum and scutum. Basicosta uniformly
black. [New Ireland] . F. bracteata

1x1 Abdomen uniformly purplish black (at most with paler reddish purple tinge on T3).

Eyes of J very nearly meeting, frons at narrowest subequal in width to anterior
ocellus and 0-03 of head-width. [Solomon Islands, ? also New Britain]
F. solomonicola

— Abdomen black with brilliant metallic transverse bands on T3-T5 ranging from
golden green to greenish copper to silvery blue-green. Eyes of ¢ slightly more
separated, frons at narrowest about one and a half times as wide as anterior
ocellus and 0-05 of head-width. [New Britain] ‘ ; : : F. heinrothi

Subgenus PSEUDOFORMOSIA Brauer & Bergenstamm

Pseudoformosia Brauer & Bergenstamm, 1889 : 126 (58). Type-species: Formosia moneta
Gerstaecker, 1860, by monotypy.

Laccura Enderlein, 1936 : 431. Type-species: Rutilia saturatissima Walker, 1861, by original
designation. Syn. n.

Diacnosis. Head partly metallic, at least on upper occiput and genal dilations. Head
of $ almost fully holoptic, upper interfrontal area eliminated by meeting of parafrontals, frons
at narrowest hardly wider than anterior ocellus. Eyes of ¢ sometimes with upper facets en-
larged. Arista short plumose, hairs much longer than basal thickness of arista. Mentum of
proboscis rather slender, tapering distally. No posthumeral setae (trace of such setae present
in occasional specimen). Usually one pair of small presutural dorsocentral setae present
(occasionally undeveloped, especially in ¢ of moneta). One post ia seta (occasional specimens
aberrant and without such seta or with trace of very small second post ia in addition). Two
sternopleural setae (1 + 1), rarely with a supernumerary third seta in addition. Postalar
callus with 4-6 setae, rather variable in degree of development. Scutum without supernumerary

32 Re Wi CROSSICE Y

setae in prescutellar region between the hindmost post acy and post dc. Scutellum not at all or
only very slightly flattened on disc. Fore coxa bare on inner half of anterior surface, except
near apex. Transverse row of erect median marginal setae of T3 present in both sexes. T5
with or without strong erect setae behind the transverse row.

DistriBuTIoNn. Molucca Islands (including Halmahera, Ternate, Batjan, Morotai,
Obi) and New Guinea. Absent from Australia and unknown (probably absent)
from the Philippines.

Discussion. This subgenus is very closely similar to Formosia s.str. but is easily
and constantly distinguishable and I consider it best to treat it as a valid subgenus.
It differs from Formosia s.str. by having two sternopleural setae and in having the
inner anterior half of the fore coxa bare (as in Euamphibolia) and by some other
more minor and less constant differences such as the presence in most specimens of a
small but distinct presutural dorsocentral seta (though this seta is most often absent
in the males of F. (P.) moneta). Pseudoformosia, like Formosia s.str., is confined to
tropical areas but has a much more restricted distribution than Formosia, as it is
apparently unrepresented anywhere west of the Moluccas (1.e. in the Oriental Region)
or in the Bismarck Archipelago or the Solomon Islands.

As known at present the subgenus is small and contains only four species; it is
not considered necessary therefore to recognize any species-groups at the present
time. However, it may be noted that the type-species, F. (P.) moneta, differs from
the other included species by having the parafrontals and epistome shining metallic
green or blue, in lacking strong setae on T5 behind the main transverse row, and in
having the upper eye facets of the male conspicuously enlarged; the other three
species (listed below) have the parafrontals and epistome non-metallic, possess some
strong setae laterally on T5 behind the transverse row, and have the male eye facets
normal. One species (pawper de Meijere) has the entire body very dark brownish
black with dark brown wings, but the other species more closely resemble typical
species of Formosva s.str. in having beautiful brilliant coppery green or bluish green
metallic patterns on the abdomen and green to violet colouring on the thorax.

Engel (1925 : 357) synonymized Pseudoformosia with Formosia and this generic
synonymy was presumably accepted by Malloch (1929 : 309), as he placed moneta—
the type-species of Pseudoformosia—in the genus Formosia. Ina slightly later paper
Malloch (1930 : 104) wrote that “The two new species [i.e. guadripunctata and
cingulata] belong to the subgenus Pseudoformosia’, though he did not say how this
subgenus differed from typical Formosia; however, Malloch’s use of Pseudoformosia
in subgeneric status means that no new status is involved in the present work.
Enderlein (1936 : 427) treated Pseudoformosia as a synonym of Formosia, but used
the latter genus in a very narrow sense (including in it only three species, most of the
species here included in Formosia s.str. being placed in Pancala Enderlein by Ender-
lein).

INCLUDED SPECIES

Formosia (Pseudoformosia) excelsa (Walker) comb. n. Motuccas. [Holo-
type examined].
F. (P.) moneta Gerstaecker. NEw GUINEA. [Holotype examined].

REVISIONARY CLASSIFICATION OF RUTILIINI 33

lucigena (Walker) syn. n. [Lectotype examined].
obscuripennis Brauer & Bergenstamm (unavailable name).
F. (P.) pauper (de Meijere) comb. n. Motuccas. [Lectotype examined].
F. (P.) saturatissima (Walker). Motuccas, ? NEw GuINEA. [Lectotype
examined].

KEY TO SPECIES OF THE SUBGENUS PSEUDOFORMOSIA

1 Thorax and abdomen unicolorous dark brownish black (sometimes slightly purplish

black) . : : . F. pauper
($ genitalia not distinguishable from saturatissima (Text- fig. 50)]

— Thorax mainly metallic green to violet-blue, abdomen with a pattern of metallic green
or cupreous spots and bands against a black background ;

2 Parafrontals metallic blue or green. Epistome metallic, distinctly shining blue or
green with violet tinges. Metallic pattern of abdominal T4 consisting of a pair of
large spots on the mid dorsum and a pair of slightly crescentic marks on the venter
against the fore margin of the tergite; T5 similarly with two metallic areas on
each side, one laterodorsal and the other ventral : : : . F. moneta

— Parafrontals thickly whitish or pale brownish pollinose, non- -metallic. Epistome not
metallic, dark brown. Metallic pattern of T4 consisting of a single continuous band
around each side from near the mid dorsum to the mid venter; T5 with a on
large metallic area on each side é 3

3 Abdominal T3 with a single broad metallic band around each side which i is con-
tinuous from near the mid dorsum to the mid venter. Wings clear hyaline or
almost so. Surstyli of J genitalia rather narrow and pointed (Text-fig. 51)

nN

F. excelsa
[This nominal species is known only from the teneral $ holotype in rather poor
condition but appears to be distinct from saturatissima]
Abdominal T3 with a pair of small submedian metallic spots on the dorsum and with
a narrow lateroventral metallic band on each side (some specimens may show traces
of metallic colour slightly interconnecting the spots and bands). Wings brown
with the infuscation most intense along the veins. nee of 3 ia ar wider and
less sharply pointed (Text-fig. 50) . ‘ : F. saturatissima

Subgenus EUAMPHIBOLIA Townsend stat. n.

| Euamphibolia Townsend, 1916: 618. Type-species: Rutilia fulvipes Guérin-Méneville, 1843

[=Rutilia speciosa Erichson, 1842], by original designation.

| Hega Enderlein, 1936 : 419, 421. Type-species: Hega viridicingens Enderlein, 1936 [=Rutilia
complicita Walker, 1861], by original designation. Syn. n.

Chromocharis Enderlein, 1936 : 420, 432. Type-species: Rutilia atribasis Walker, 1861, by
original designation. Syn. n.

| Dracnosis. Head thickly pollinose, nowhere metallic, at most only slightly shining in some
‘lights on upper occiput. Eyes of J not very strongly approximated, interfrontal area always
distinguishable on upper part and at least as wide as parafrontal, frons at narrowest subequal
‘in width to or much wider than facial carina. Upper facets of 3 eyes not or only slightly en-

‘larged. Arista micropubescent, pubescence at most only as long as basal diameter of arista.

' Proboscis in profile not noticeably tapering towards the apex, upper and lower edges subparallel.

_Posthumeral setae present, at least one each side and often more. 2-5 presutural dorsocentral

| setae (sometimes weak and scarcely differentiated from the hair ing). 2-3 post ia setae (some-
times small fourth seta and g sometimes with only one main fost ia seta preceded by a very
small fine inconspicuous setula weakly differentiated). Two sternopleural setae (1 + 1),

34 R. W. CROSSKEY

anterior stp/ sometimes very weak and hair-like ing. Postalar callus with three setae. Scutum
without supernumerary prescutellar setae between the hindmost acy and dc setae. Disc of
scutellum not conspicuously flattened. Fore coxa bare on inner half of anterior surface (except
near apex). 73 with a transverse row of median marginal setae either in both sexes or in 9
only. T5 with long dense hair behind the transverse row of setae in which some long strong
setae developed.

DistTrisuTion. From the Molucca Islands (including Halmahera, Ternate,
Batjan, Seram, Buru, Run Island) through New Guinea to the Solomons, and in
Australia from Queensland to Tasmania, also Western Australia. [No specimens
have been seen from Bismarck Archipelago or Aru Islands but these areas are almost
certainly within the subgeneric range: Osten Sacken (1881) recorded a specimen of
pretiosa Snellen van Vollenhoven from Wokan in the Aru Islands. ]

Discussion. Townsend (1916) proposed the genus EuampMbolia for Rutilia
fulvipes Guérin-Méneville, a large and attractive Rutiliine with a black-and-white
body pattern superficially similar to that of some Amphibolia species (Brauer &
Bergenstamm, 1889 : 418, had in fact placed fulvipes, though attributing the species
to Macquart, in the genus Amphibolia). This distinctive species was described by
Erichson (1842) with the name Rwtilia speciosa in the year before Guérin-Méneville’s
(1843) description of fulvipes was published; nevertheless it is clear from Guérin-
Méneville’s account that he knew of Erichson’s name, for he expressed some doubt
as to whether his fulvipes was really distinct from Erichson’s speciosa. Later authors
have had no doubts that the names are synonyms: the synonymy of fulvipes with
speciosa was implied by Malloch (1927 : 351) when he cited speciosa as the genotype of
Euamphbolia, was formally established by Townsend (1932 : 38), and was cited by
Enderlein (1936 : 430) and Townsend (1938 : 414). Direct comparison of original
types is not possible, for although Erichson’s types still exist (and have been
examined) that of Guérin-Méneville is lost: but Guérin-Méneville’s detailed descrip-
tion of fulvipes applies so perfectly to Erichson’s types of speciosa that I here accept
the synonymy as certainly correct (as this is in accord with previous practice there
is no need of neotype designation for fulvipes). (It should be noted that Townsend,
1932, 1938, was in error to state that a holotype—sex unspecified by Townsend—of
fulvipes from New South Wales is in Paris Museum: there is no such specimen in
Paris, although Macquart’s collection contains material determined as fulvipes by
Macquart.)

Malloch (1927 : 351) wrote of speciosa as follows: ‘This is the genotype of Euampiu-
bolia Townsend, but I do not consider it entitled to generic separation from Formosia,
the only character distinguishing it being the pubescent arista. It may ultimately be
accorded subgeneric rank with atribasis.’ I completely agree with Malloch’s views
(although there are certainly more characters than the pubescent arista which
distinguish Euamphibolia from Formosia s.str.) and here accord to EuampMboha
the status of a subgenus within Formosia s.1. In agreement also with Malloch, I
consider Walker’s airibasis to be consubgeneric with speciosa, and the generic name
Chromocharis Enderlein (of which atribasis is type-species) therefore enters into new
synonymy with Euamphibolia.

The generic name Hega Enderlein is also a new synonym of Euamphibaea

7

REVISIONARY CLASSIFICATION OF RUTILIINI 35

Enderlein characterized Hega, and differentiated it from related genera, by the lack
of median marginal setae on abdominal T3, without realizing that in the group of
species to which the type-species (vividicingens) belongs the lack of median marginals
on T3 is only a sexual character. The males lack such setae and the females possess
them. The genitalia of the males, with the very heavy subquadrate surstyli, are
of an identical kind with the male genitalia of speciosa, and confirm beyond any
doubt that Hega is a synonym of Euamphibolia.

The subgenus Evamp/ibolia as here re-defined is easily distinguished from the
other subgenera of Formosia by the characters given in the accompanying sub-
generic key. Apart from the characteristic shape of the $ surstyli (see Text-figs
40, 52, 53), Euampmbolia is immediately and most easily distinguished from
Pseudoformosia and Formosia s.str. by the presence of only three setae on the postalar
callus and by the rather broad non-tapering shape of the mentum when seen in
profile (Text-fig. 12).

At present Euampbolia contains only a small number of species. Two of these,
speciosa and fusca sp. n., are very distinct from the others because of their coloration,
but as there are so few species and no really tangible structural characters on which
they can be grouped it has not been thought useful to recognize any species-groups
as formally defined entities.

Very little is known of the early stages and host relations. Townsend (1936 : 152)
has briefly described the first stage larva, and has also figured the egg and first stage
larva under the name E. fulvipes (Townsend, 1942, plate 21, fig. 147; plate 30, fig.
249; and plate 43, fig. 250). Two specimens have been seen of an undetermined
species of the subgenus from Lae, in New Guinea, which have associated puparia and
data showing that the larvae were found in a sago-palm trunk parasitizing the larvae
of a Cetoniid beetle. Otherwise nothing is yet known.

INCLUDED SPECIES

Formosia (Euamphibolia) atribasis (Walker). Motuccas. [Lectotype exa-
mined].
F. (E.) complicita (Walker) comb. n. Motuccas; West Irtan. [Holotype
examined].
pretiosa (Snellen van Vollenhoven) syn. n. [Lectotype examined].
sapphirina (Walker) syn. n. [Holotype examined].
smaragdifera Bigot syn. n. [Lectotype examined].
viridicingens (Enderlein) syn. n. [Holotype examined].
F. (E.) engeli (Enderlein) comb.n. RuNIsLAND, Buru. [Lectotype examined].
F. (E.) faceta (Enderlein) comb. n. AvsTRALIA (Queensland); NEw GUINEA.
[Holotype examined].
F. (E.) fusca Crosskey sp. n. SERAM. [Holotype examined].
F. (E.) smaragdina Malloch. AvusTRALIA (Queensland). [Holotype examined].
F. (E.) speciosa (Erichson). AvusTRALIA (Australian Capital Territory, New
South Wales, Queensland, Tasmania, Victoria, Western Australia). [Lecto-
type examined].
fulvipes (Guérin-Méneville) [Holotype lost}.

36

. Wi. CROSSIKE Y

KEy TO SPECIES OF THE SUBGENUS EUAMPHIBOLIA

Legs entirely reddish yellow. Dorsum of thorax black with bold white spots and
vittae. Abdomen with black-and-white pattern. Palpi yellow. Pleural regions
of thorax with yellow to golden orange hair. Hair of coxae and femora largely
golden orange. [Australia and Tasmania] ‘ : . F. speciosa
Legs entirely black or brownish black. Dorsum of oes ae so, eae to blue-
violet or if blackish (fusca) then without white spots. Abdomen not black-and-
white. Palpi dark brown or blackish. All thoracic hair black. Hair of coxae
and femora all black . : ; : : c : : 2
Thorax and abdomen unicolorous Gas plc prone [Seram] F. fusca sp. n. (p. 37)
Not unicolorous dark species, dorsum of thorax and usually also much of the abdomen
metallic greenish, cupreous, blue or violaceous, the abdomen always noticeably
banded or patterned. 3
Hair of occiput and Sostbaecee all BEeee Head pollinosity distinctly bicolorous,
white on lower parts of parafrontals and on parafacials, postorbits, and most of
genae, and pale golden brown on upper parts of parafrontals and on occiput, facial
carina, antennal foveae and epistome (golden brown colour of parafrontals best
seen from above). Thorax without white pollinosity, with only merest trace of
yellowish brown pollinosity thinly over humeral calli and fore margin of prescutum.
Dorsum of T5 with a pair of widely separated golden green subtriangular metallic
areas which extend back only about as far as the main transverse row of setae.
[Batjan] - : F. atribasis
Hair of occiput rete or ellen ini of pastbdcase neal ae ie but occasionally
brownish. Head pollinosity unicolorous white, greyish white or yellowish white.
Thorax usually with some white pollinosity on prescutum and humeral calli, often
also on mesopleuron and sternopleuron. Dorsum of T5 with metallic areas
not of thisform . 4
Dorsum of abdomen with pon Goiel eoppery na or eae bands on each side Gf
T3-15 which are clearly separated medially on all tergites (T3, therefore, as well as
other tergites with distinct black median vitta); the dark colour of the metallic
bands not strongly contrasting with black areas of tergites and abdomen appearing
generally dark. Wings very conspicuously brown. [Run Island and Buru]
F. engeli
Dorsum of abdomen with a bold pattern of golden green, reddish green, emerald or
blue metallic areas on a black background, the abdomen not appearing rather
uniformly dark. Wings mainly clear hyaline or nearly so (except for usual
black basal area), only weakly suffused with yellowish brown along the veins : 5
Abdominal T5 with the metallic areas covering most of the dorsum, at most only the
posterolateral corners of the tergite and a very fine inconspicuous median line
black. 9 without definite sternopleural pollinose spot and with rather thin white
pollinosity over mesopleuron which has a distinctly shifting appearance from differ-
ent angles. Upper occiput metallic greenish adjacent to the postocular setulae. ¢
frons at narrowest 0:14-0:15 of head-width. [Moluccas and West Irian]
F. complicita
Abdominal T5 with metallic areas confined to middle part of tergite dorsum, some-
times evanescent so that T5 is virtually all black in J, metallic areas of T5 dorsum
usually in form of two irregular elongate golden green areas which nearly meet
posteriorly in the tergite depression and together are roughly V-shaped. 2 with
a small but definite pollinose spot on the upper sternopleuron and with a boldly
marked white pollinose spot on mesopleuron which does not strongly shift with the
light direction. Upper occiput either metallic or non-metallic adjacent to postocu-
lar setulae. ¢ frons at narrowest 0-12—0-13 of head-width : 6
Upper occiput evenly white pollinose and showing little or no trace of meee icoleee

REVISIONARY CLASSIFICATION OF RUTILIINI 37

near the postocular setulae. White pollinosity of humeral calli and prescutum of 2
thick and very conspicuous to naked eye, making anterior half of thoracic dorsum
appear rather greyish (except for the black vittae). Pollinose spot of sterno-
pleuron weakly marked and partly brownish. Submedian pair of black marks on
the anterior edge of scutum (i.e. the ends of the inner pair of black mesonotal
vittae) more or less connected by dark brown coloured area medially. Small
species, length 12.5.16 mm. [Queensland and New Guinea] ; : . FF. faceta
— Upper occiput distinctly metallic green in the area of the postocular setulae. White
pollinosity of humeral calli and prescutum of 9 inconspicuous to naked eye, not
distinctly reaching to transverse suture, whole mesonotum appearing bright
metallic green or blue-green. Pollinose spot of sternopleuron very distinct, almost
entirely white like the mesopleural spot. Submedian pair of black marks on
anterior edge of scutum (ends of the inner pair of black mesonotal vittae) small
and square and widely separated by the metallic green colour, which reaches
medially to the front edge of the scutum (transverse suture). Large species, length
17°5-20 mm. [Queensland] : F. smaragdina

Formosia (Euamphibolia) fusca sp. n.
(Text-fig. 40)

6- Head. Ground colour blackish brown, slightly more reddish brown on facial carina,
antennal foveae and upper genae; lunula shining yellowish brown, interfrontal area velvety
brownish black; head rather thickly yellowish white pollinose on anterior parts, more thinly
greyish white pollinose on occipital regions. Parafrontal and genal hair black, occipital hair
reddish brown. Eyes not very strongly approximated and upper facets not enlarged; upper
interfrontal area distinct and nearly twice as wide as parafrontal, frons at narrowest about
| 0-14 of head width and wider than facial carina. Ocelli slightly raised, ocellar setae undeveloped.
_ Facial carina nearly twice as long as epistome, subparallel-sided. Gena one-third (0-33) of eye
jheight. Parafacial about 3-5 times as wide as width of third antennal segment and much
wider than length of this segment. Antennae dark reddish-brown, very small and falling short
/ of mouth-margin by very much more than their own length; third segment about 2-1 times as
| long as second segment; arista micropubescent. Palpi brown with paler tips. Thorax. Black-
/ish brown with very thin brownish pollinosity, non-metallic; scutellum slightly more reddish
/brown than scutum. Thoracic hair entirely black. Chaetotaxy normal for subgenus but
| setae rather small [post ia setae and postalar setae of left side and some marginal scutellar setae
/missing from holotype, but pores distinct]. Anterior sternopleural seta minute, almost hair-
like. Scutellum with five pairs of marginal setae, and one or two smaller irregular submarginals.
Wings. Basicostae and basal spots extremely dark, almost black; membrane mainly hyaline
_except for narrow traces of yellowish brown colour immediately adjacent to veins. Calyptrae
.dark brown with brown marginal hair. Legs. Black with entirely black hair; femora not
conspicuously attenuate. Mid tibia without ad setae and without definite p setae; hind tibia

without pd setae. Claws very long. Abdomen. Entirely blackish brown with black hair,
_concolorous with thorax. Dorsal hair exceedingly fine and abundant, recumbent on T3 and
| very dense medially, semi-erect on T4 and erect on T5. T3 without median marginal setae;
| T5 with some long setae developed among the long fine hair posteriorly to the transverse row
}of setae. Depression of T5 well formed and conspicuous. Genitalia as in Text-fig. 40. Mea-
|suvements. Body length 18-2 mm, wing length 18-5 mm [one specimen, holotype].

9. Extremely similar to § except for usual sexual differences. Interfrontal area only
' slightly narrowing dorsally; proclinate orbital setae absent. Vertex 0:25 of head width. Facial
carina more pinched in at lower end than in gj. Body length 20-0 mm, wing length 18-8 mm
[one specimen].

38 RoW. CROSSKE ¥

MATERIAL EXAMINED
Holotype g, InponEs1A: Moluccas, Seram (Ceram), Mansela, 2500 ft, 1919
(Pratt). In British Museum (Natural History), London. [Genitalia on slide.]

Paratype. 1 9, INnDoNESIA: Moluccas, Seram (A. R. Wallace) (BMNH, London).
DISTRIBUTION. Known only from the island of Seram (Ceram) in the Moluccas.

AFFINITIES. A very distinctive species readily identified at a glance by the
unicolorous blackish brown colour and large size; no particular affinity with another
species is evident at present. Although only one male and one female are available
so far, I have no hesitation in describing the new species because it is so clearly
distinct from all other known species of the subgenus Euamphibolia, to which fusca
sp. n. is certainly assignable.

Genus FORMODEXIA Gen. n.
Type-species: Rutilia volucelloides Walker, 1861.

Diacnosis. Facial carina large, with subparallel sides and at most only slightly bulbous
medially, convex on outer surface. Epistome not very strongly prominent, weakly set off from
carina by shallow depression. Eyes of very strongly approximated so that upper frons slightly
narrower than facial carina, the upper facets not noticeably enlarged. Parafacials bare. Buc-
cal opening very narrow, in ¢ hardly wider than facial carina. Genal dilation moderately
developed, not reaching forward as far as front level of eye. Head nowhere metallic. Arista
pubescent. Palpi of both sexes exceptionally long and slender. Proboscis rather long and
slender and slightly tapering before the labellae. Prosternum and prosternal membrane bare.
Scutellum with apical pair of setae inserted at lower level than other marginal setae; total of
six or seven pairs of marginals; disc of scutellum not flattened. Postalar callus with 5 setae.
Postalar wall with dense hair tuft. Suprasquamal ridge bare. Upper calypter normal.
Tegula without the usual long wiry posterior setulae. Costal base explanate and with well
formed close-set curved marginal fringe, wings appearing to have basal ‘shoulder’ to naked eye.
Abdomen without downwardly-directed spiniform setae on ventral margins of tergites; T3
without either median or lateral marginal setae; intermediate tergites without discal setae.
T5 convex, without median depression or prominent posterolateral corners, apical part (behind
the single transverse setal row) sharply bent downwards in ¢.

DISTRIBUTION. Only known from the Moluccas, including the islands of Halma-
hera and Batjan; probably occurring also in other islands of the Moluccas group.

Discussion. The curious Rutiliine species described by Walker as Rutilia
volucelloides (of which two other Walker names are synonyms, as indicated later)
from the Molucca islands has a suite of characters which preclude it from being
placed satisfactorily in any of the genus-group segregates of Rutiliini already des-
cribed, and in order to present a balanced classification of this tribe it is necessary
to assign volucelloides to a new genus, for which the name Formodexia gen. n. is
proposed.

Formodexia gen. n. agrees with Formosza, and differs from all other Rutiliine genera,
by having the postalar wall haired, but it differs from Formosia in having the
apical scutellar setae set lower than the others, in the explanate costal base and
lack of wiry posterior hairs on the tegula, and in lacking the strong downward-point-
ing spinous setae on the abdominal venter which are characteristic of Formosia;

REVISIONARY CLASSIFICATION OF RUTILIINI 39

the same differences also distinguish Formodexia from Rutilodexia. Formodexia

resembles Rutilia in having the apical scutellar setae at a lower level than the other

marginal setae, but is easily separated from all Rutilia species by the hairy postalar
wall and from almost all Rutilia by having the suprasquamal ridge bare.

There is a strong superficial resemblance between Formodexia and Prodiaphania,
especially because of the explanate costal bases of both sexes in both genera and
because of the strongly narrowed buccal opening, but Formodexia is very easily
distinguished from Prodiaphania by the exceedingly long slender palpi (palpi ex-
tremely reduced in Prodiaphania), by the haired postalar wall (postalar wall bare in
Prodiaphania), by the pubescent arista (arista plumose in Prodiaphania) and by the
normal small upper calypter (upper calypter much enlarged in Prodiaphania).

Although Formodexia is unlikely to be confused with the remaining Rutiliine
genera it may be useful to note the following distinctions: the new genus differs from
Chrysopasta by having the postalar wall haired and the suprasquamal ridge bare and
in the bare parafacials, and the same features of the postalar wall and the supra-
squamal ridge distinguish it also from Amphibolia (but some species of the latter
agree with Formodexia in having bare parafacials). From Chetogaster, which only
doubtfully belongs to the Rutiliini, it may be separated at once by the five setae on
the postalar callus (only two such setae in Chetogaster).

Additional features of Formodexia to those cited in the diagnosis, given to facilitate
comparison with various subgenera of Rutilia and Formosia, are as follows: no
development of strong spiniform setae on either thorax or abdomen; setae of pre-
scutum and scutum reduced, post ia setae absent; two sternopleural setae (1 + 1);
_ claws of unusually small; 2 without proclinate orbital setae; hind tibia with short
well developed and close-set ad fringe; head of $ appearing unusually small in
' relation to body size; both sexes with a row of well developed setae along the basal
| half of the a surface of the mid femur.

_ Asurprising aspect of Formodexia is the extreme paucity of material so far known
of volucelloides, the only species: I know of only five specimens, all in the British
Museum (Natural History), of which three are Walker’s types of three synonymic

names and the other two are old specimens from Batjan that formed part of Bigot’s

collection. Of the five specimens, only the primary type (lectotype herein desig-
nated) of volucelloides is a male, but the species is so distinctive that there is no doubt

_ whatever that the four female specimens are conspecific with this male; as a conse-
quence, Walker’s names ¢tvixoides and ignobilis fall as new synonyms of volucelloides.

Enderlein (1936), without seeing Walker’s types and therefore without any real
idea of their characters, assigned ignobilis to Chrysorutilia Townsend and assigned

_ volucellordes and trixoides to his genus Donovanius, thus in effect associating these

names with Rutilia (for Chrysorutilia and Donovanius are, in the present treatment,

regarded as not generically distinguishable from Rutilia). As already shown, all

_ three specific names belong to one species which is not assignable to Rutilia.

' Attention may usefully be drawn here to an error in Walker’s citation of type-

locality for ignobilis: this was originally recorded in error as Gorrite (in Brazil),

but the correct type-locality (as indicated by a label on the specimen in Walker’s

_ writing) is southern Gilolo (= Halmahera) in the Molucca islands.

40 RW. CROSSIKE ¥

INCLUDED SPECIES

Formodexia volucelloides (Walker) comb. n. Motuccas. [Lectotype exa-
mined].
trixoides (Walker) syn. n. [Holotype examined].
ignobilis (Walker) syn. n. [Holotype examined].

Genus RUTILODEXIA Townsend

Rutilodexia Townsend, 1915 : 23. Type-species: Rutilia angustipennis Walker, 1859, by original
designation.

Bothrostiva Enderlein, 1936 : 413. Type-species: Bothrostiva prisca Enderlein, 1936, by original
designation. Syn. n.

Rutilosia Paramonov, 1968 : 355. [Name published in footnote to generic key: unavailable
under Article 13 (b) of International Code of Zoological Nomenclature, no fixation of type-
species. ]

Diacnosis. Facial carina large and subparallel-sided, at most slightly widened medially,
often slightly sulcate. Epistome not strongly prominent, separated from facial carina by very
shallow depression only. Eyes of § not very strongly approximated, upper frons at least as
wide as and usually a little wider than facial carina, upper facets not enlarged. Parafacials
bare. Buccal opening wide, very much wider than facial carina in both sexes. Genal dilation
weakly developed, anterior edge very remote from epistome. Head nowhere metallic.
Arista pubescent. Palpi fully developed normal size. Proboscis short, rather slender and
tapering before labellae. Prosternum and prosternal membrane bare. Scutellum with apical
pair of setae inserted at lower level than other marginal setae (apicals sometimes hair-like) ;
total of 7-11 pairs of marginals; disc of scutellum not noticeably flattened. Postalar callus
with 4-6 setae. Postalar wall bare (at most a very few hairs adjacent to those on callus edge).
Suprasquamal ridge bare. Upper calypter normal. Tegula with normal long wiry posterior
setula. Costal base not explanate. Abdomen with slightly spiniform setae ventrally on tergite
margins which are directed downwards and backwards; T3 without median marginal setae in g,
with single pair of such setae in 2; intermediate tergites without discal setae. T5 broad, without
or with at most only slight median depression, posterior corners not prominent.

DISTRIBUTION. Aru Islands, New Guinea and New Britain.

Discussion. It is considered best to recognize Rutilodexia Townsend as a valid
genus for a small group of species found (so far) only in the Papuan subregion and
differing on total suite of characters from the species placed in Rutilia s.1., though
undoubtedly Rutilodexia is very closely allied to Rutilia and does not differ from it
by very convincing characters. The general appearance of Rutilodexia species is
very similar to some Rutilia (especially in the subgenus Donovanius Enderlein), and
Rutilodexia has the bare postalar wall and low-set apical scutellar setae exactly as in
Rutilia; furthermore, the facial carina is exactly of the Rutilia type. The supra-
squamal ridge is, however, bare in Rutilodexia and this character provides a distinc-
tion from almost all of Rutilia species; in Rutilia the suprasquamal ridge is nearly
always haired, but there are some species which (on overall balance of characters)
must be placed in Rutilia even though the ridge is bare. The best feature for
distinguishing all Rutilodexia from all Rwutilia seems to lie in the genal dilation
(Text-figs 4 & 5); in Rutilodexia this is unusually small and its anterior limit is formed
by an oblique ridge extending downwards from behind the eye; in Rutilia the genal

REVISIONARY CLASSIFICATION OF RUTILIINI 41

dilation is more or much more extensive and rounded anteriorly. In Rutilodexia
the anterior part of the genal dilation is very remote from the epistome and is not
as far forward on the head as the level of the front of the eye; in Rutilia the anterior
part of the dilation normally reaches forward to or beyond the level of the front of
the eye and is thus less remote from the epistome.

The ¢ genitalia of Rutilodexia species have no special characteristics; the surstyli
and cerci have a simple form (Text-fig. 90) generally resembling that found in a
number of Ruwutilia s.1. species.

The form of the genal dilation and close resemblances in all other characters
indicate without doubt that Bothrostira prisca Enderlein is congeneric with, though
a separate species from, Rutilodexia angustipennis (Walker), and Enderlein’s generic
name Bothrostiva therefore falls in new synonymy with Rutilodexia. It is certain
also that Paramonov’s (1968) posthumously published and unavailable name
Rutilosia applies to the same concept (because the manuscript notes left at his
death show that Paramonov included angustipennis Walker in Rutilosia without
him appreciating that this nominal species was already type-species of Rutilodexia).
Comparison of Enderlein’s types of B. prisca ($) and Idania ralumensis (2), in con-
junction with later collecting of both sexes, shows that both names apply to the same
species; as valumensis is certainly the female of prisca (though Enderlein (1936 :
409, 413) had the nominal species in separate genera) the names are here placed in
new synonymy, with prisca treated as the valid name because based on a male type.
R. papua (Bigot) is perhaps the same species as R. angustipennis (Walker) but
there is not sufficient evidence at present to justify synonymy.

Nothing is known of the hosts of Rutilodexia species.

INCLUDED SPECIES
Rutilodexia angustipennis (Walker). Aru IsLtanps. [Holotype examined].
R. papua (Bigot) comb. n. NEw GuiNnEA. [Lectotype examined].
R. prisca (Enderlein) comb. n. NEw Britain. [Holotype examined].
valumensis (Enderlein) syn. n. [Lectotype examined].

Kry TO SPECIES OF THE GENUS RUTILODEXIA

[Note : The primary types of angustipennis and papua are male and female respectively, and
the opposite sexes cannot yet be positively associated. |

© Males _. : : ‘ é : : : : ; : : : : 2
— Females . : 4
2 Hind tibia with a strong submedian ad seta and a very realli inconspicuous ad fringe.
Hind femur with a row of strong av setae on the apical half. [Aru Islands & New
Guinea] : : R. angustipennis

— Hind tibia without a submiedian ad seta and - Ww ith 2 a well dev dloped regular close-set ad
fringe. Hind femur without strong av setae on apical half, the vestiture hair-like
(New Guinea & New Britain] :

3 Dorsum of thorax and abdomen contrasting in colour; the es and scutellum
metallic dark green or bluish green and the abdomen tawny yellowish with a
conspicuous black median line. [New Britain] 4 ; . &. prisca

— Dorsum of thorax and abdomen more or less concolorous, dark brow nish or blackish
brown with a purplish green or violaceous sheen and without a noticeable dark
median line on the abdomen. [New Guinea] . : Undetermined sp. (? papua 3)

42 RoW. CROSSKEY

4 Hair of the barette, posteroventral parts of the pteropleuron and outer edge of
postalar callus golden yellow (in contrast with black hair on remainder of thorax).
Prescutum with only indistinct traces of dark vittae. Head tawny or dark reddish
with brassy yellow pollinosity. Wings rather conspicuously smoky on apical half
and yellowish on basal half. [New Britain] . . &. prisca

— Hair of barette and all hair of pteropleura and postalar ‘calli pad like the rest of the
thoracic hair. Prescutum with four bold black-brown vittae. Head blackish
brown with pale greyish yellow age ee not seu Er yellow and
smoky brownish. [New Guinea] . ; : . RR. papua

Genus RUTILIA Robineau-Desvoidy

Rutilia Robineau-Desvoidy, 1830 : 319. Type-species: Tachina vivipava Fabricius, 1805, by
subsequent designation of Crosskey (1967 : 26).

Diacnosis. Facial carina large, often broad with subparallel sides, sometimes slightly
knob-like or slightly evanescent ventrally. Epistome slightly to very strongly prominent.
3 head never holoptic, upper frons at least as wide as facial carina, upper eye facets not enlarged.
Parafacials bare or haired. Buccal opening normal, wider than facial carina in both sexes.
Genal dilation well developed, usually rounded anteriorly and extending forwards nearly to
level or beyond level of front of eye. Head normally mainly pollinose, partially metallic in
some forms. Arista virtually bare to very short-plumose, usually pubescent. Palpi fully
developed normal size (except reduced in micropalpis). Proboscis short with sides subparallel
in profile or at most only a little tapering before labellae. Prosternum and prosternal membrane
bare or haired. Scutellum with apical pair of setae inserted at lower level than other marginal
setae (very rarely apicals absent); total of 3-10 pairs of marginals; disc of scutellum convex
or flattened. Postalar callus with 3—5 (rarely 6) setae. Postalar wall bare (at most a very few
hairs adjacent to those on callus edge). Suprasquamal ridge almost always haired, bare in a few
species. Upper calypter normal. Tegula with normal long wiry posterior setulae. Costal
base sometimes distinctly explanate. Abdomen with marginal vestiture of tergite venters
weak and semi-recumbent (directed backwards), if bristling slightly spiniform then not directed
vertically downwards; T3 with or without median marginal setae; intermediate tergites without
discal setae. T5 of varied form, convex truncate conical to broad and flattened with median
depression.

DISTRIBUTION. Especially well represented and widespread in Australia, but
occurring also in Oriental Region from India and Ceylon through Malaya to Philip-
pines and Timor; poorly represented in New Guinea, occurring in Solomon Islands,
New Hebrides, Fiji, Samoa and Lord Howe Island. Absent on present evidence
from Moluccas and Bismarck islands.

Discussion. The genus Rutilia in the wide sense here adopted (which corres-
ponds in the main with the sense of the genus adopted by Malloch and Paramonov)
is the largest genus of Rutiliini and contains nearly half the described species. It is
the dominant element in the Rutiliine fauna of Australia, and contains most of the
large metallic and boldly patterned species that are such a conspicuous element in
the Australian dipterous fauna. The genus is not easy to define in a completely
satisfactory way, and the diagnosis has perforce to take into account several species
that differ from typical Rutilia in some conspicuous feature, even though on their
totality of characters they must clearly be assigned to the genus: for example, some
species have the suprasquamal ridge bare (e.g. cungulata Malloch and confusa Malloch,
which were erroneously placed in Formosia by Malloch because of this), and

REVISIONARY CLASSIFICATION OF RUTILIINI 43

occasionally specimens occur which lack the apical setae of the scutellum. The
differences between Rutilia and other Rutiliine genera have been alluded to else-
where under the different genera and need not be repeated here, but it might be
useful to emphasize the chief differential characters which separate Rwutilia and
Formosia, for there has always been some difficulty in separating these genera
satisfactorily since the time when Guérin-Méneville (1843) first distinguished them.
The table below indicates the main differences:

Rutilia

Postalar wall bare (at most a very few
hairs immediately below edge).

Suprasquamal ridge haired (bare in a
few exceptions).

Apical scutellar setae inserted below
level of other marginal setae (very
rarely absent).

Marginal setae of abdominal tergite

Formosia
Postalar wall with dense hair tuft.

Suprasquamal ridge bare.

Apical scutellar setae inserted level with
other marginal setae.

Marginal setae of abdominal tergite

venters usually very weak, recum-
bent, at most only slightly spiniform
and directed backwards as well as
downwards.

venters very strong, spiniform, direct-
ed straight downwards.

Some supposed genera have been split off from Rutilia by various authors but not
generally accepted as separate genera (e.g. Microrutilia Townsend, Chrysorutilia
Townsend), and Enderlein (1936) fragmented Rutilia into eleven genera placed in
two subtribes, using such features as the present or absence of median marginal
setae on abdominal T3 or the presence or absence of a median depression in the last
abdominal tergite (T5) as generic characters. In the present work none of these
genera has been considered valid as a taxon worthy of generic rank, but a few have
been redefined in the light of the characters shown by the type-species and recog-
nized as subgenera within Rutilia; in all, seven subgenera are here recognized and
defined, for six of which there are pre-existing genus-group names available (these
including Rutilia s.str.) and for one of which a new subgeneric name is proposed
(there being no previously published name applying to the taxon concerned). The
subgeneric name Neorutilia Malloch and the following sixteen generic names are
synonyms of Rutilia s.l. (the correct synonymic distribution of these names according
to the subgeneric classification here proposed is shown at the head of each subgenus) :
Agalmia Enderlein, Chrysorutilia Townsend, Donovanius Enderlein, Eucompsa
Enderlein, Grapholostylum Macquart, Habrota Enderlein, Idania Enderlein, Mene-
villea Enderlein, Microrutilia Townsend, Philippoformosia Townsend, Prosenostoma
Townsend, Psaronia Enderlein, Psaroniella Enderlein, Pogonagalmia Enderlein,
Stiraulax Enderlein and Zoramsceus Enderlein.

Though the numerous species of Rutilia s.1. (some 60 species at present, but several
undescribed species known) show great diversity in their general appearance—size,
colouring, pollinose patterns—and though a few species are difficult to place satis-

44 Rk. W. CROSSKE Y

factorily in any particular segregate, it is generally true that the great majority of
species can be placed alongside obviously allied species where together they form
natural subdivisions of the genus, each of which can be reasonably well charac-
terized and differentiated from other such segregates. It is here considered best to
treat these major groupings of the species as subgenera, a course which has not
previously been adopted with Rutilia (except for Malloch’s (1936) recognition of his
new species simplex as representing a new subgenus, and his recognition of subgenus
Microrutilia in the same paper). The seven subgenera recognized can be distin-
guished by the accompanying key and diagnoses, but it may be useful to draw
attention here to some general conclusions to which I have come before formulating
these subgenera.

It is certainly unwise in Rutilia (as indeed generally in the Tachinidae) to aggre-
gate species into a defined taxon merely on the common possession of a single
character, or to attach too great an importance to some single striking attribute that
may well have, and usually almost certainly has, been evolved more than once within
the whole complex of forms. This course forced Malloch to place species such as
confusa Malloch and cingulata Malloch out of Rutilia and in Formosia instead,
simply because on the single character of bare suprasquamal ridge they are atypical
for Rutilia, though on their overall totality of characters these species clearly fit
Rutila and not Formosia. Yet within Rutilia the rest of the characters shown by
confusa and cingulata leave little doubt that these two species are not closely allied,
and an unnatural group would be formed by defining a taxon (subgenus or species-
group) which brought these species together: it seems certain that bareness of the
suprasquamal ridge has arisen more than once within Rutilia, so that while the charac-
ter can be usefully used as one of the characters of an infrageneric taxon it cannot
be used as an exclusive diagnostic feature. Similarly, the depression in the last
visible abdominal tergite (T5), which is such a conspicuous feature of many Rutilia
species and was used as a generic character by Enderlein, has undoubtedly in my
view arisen at least twice in separate lines of evolutionary development in the
genus: it occurs principally in the subgenus Donovanius but is also found in Rutilia
s.str., groups which seem to resemble each other convergently but differ much when
all their characters are taken into account.

In Rutilia the chaetotaxy is often very variable, the number of setae and their
degree of development often showing great inconstancy within a species and between
sexes (females normally have a stronger chaetotaxy and may show development of
certain setae that are usually totally lacking in males), and also sometimes showing
imperfect bilateral symmetry. Hence chaetotactic characters must be used with
great caution and long series of specimens require study before any conclusions can
be made about which setae, if any, are more or less dependable. The acrostichal,
dorsocentral, intra-alar and supra-alar setae are, speaking generally, too inconstant
in number and strength to be of real value in providing group characters or specific
characters, and the sternopleural and posthumeral setae (especially the latter) show
some variability which makes their usefulness limited (though the sternopleurals can
usefully be used, and are here used, as a supporting character in defining the sub-
genera). The humeral callus has two moderately strong setae on the outer half in

REVISIONARY CLASSIFICATION OF RUTILIINI 45

nearly all specimens, and some Ruwtilia s.l. have one or two very fine humerals
developed and distinguishable from the hair on the inner half of the callus as well;
though a somewhat intangible character, it does seem that the setae on the inner
half of the callus are almost never developed in the subgenus Chrysorutilia, whereas
at least one and usually two humerals are (even if very fine) normally differentiated
from the hair on the inner half of the callus in other subgenera.

An extremely important chaetotactic character among Ruitilia s.l. is the number
of strong setae on the postalar callus, and I find this to be one of the most useful
characters that exists (on present knowledge) for dependably segregating the species
into their respective subgenera. This feature of the chaetotaxy, whether there are
three postalar setae or whether there are four (or more) such setae, provides an almost
completely dependable character for distinguishing some subgenera from others—
though very occasional specimens are found in which there are three postalars on
one side and four on the other. If the species of Rutilia are assembled on the basis
of their common possession of three and four postalar setae it is found that the
resulting groupings appear without doubt to reflect a natural dichotomy within the
genus, and that distinct taxa (subgenera) can then be defined on the basis of several
other characters taken in combination with the postalar character. Omitting the
small subgenera Neorutilia and Ameniamima subgen. n. (of which the affinities are
rather obscure) the great bulk of Rutilia species are comprised here in the present
re-classification in five subgenera, three in the section possessing the three postalar
setae and two in the section with four such setae. The subgenera with three
postalars (Rutilia s.str., Microrutia and Grapholostylum) have a well developed
anterior sternopleural seta, lack a hind tibial fringe, and have the hair of the supra-
squamal ridge rather short and sparse, and are considered very closely allied; the
subgenera with four postalars (or more) (Donovanius and Chrysorutilia) seem
similarly close to each other (although their ¢ genitalia differ strikingly) and comprise
forms without definite anterior sternopleural setae, with a well developed hind tibial
fringe, and with extremely long thick bushy and often crinkled hair on the supra-
squamal ridge. Distinctions between the seven subgenera recognized are em-
phasized in more detail in the treatment of each subgenus.

There is a strong tendency for the constituent species within each subgenus to
have the same or similar type of colour pattern, though species may look super-
ficially rather different because of different intensities of colour and of widely
differing sizes. In Rutilias.str. the predominant coloration is brown with little or no
development of metallic colouring and no spot patterns, whereas in Chrysorutilia
the species are rather homogeneously metallic green to blue-violet, with yellow heads
and some coppery or blackish patterning or iridescence ; in Amenitamima the colouring
is green to purplish black, and both thorax and abdomen have a pattern formed of
boldly marked white pollinose spots. A general indication of the range of colour
and pattern, together with size, found in each subgenus has been given as an appendix
to each of the subgeneric diagnoses.

The male genitalia and the form of the 5th (subgenital) abdominal sternite of
the male provide some characters of taxonomic value at subgeneric and specific
| level, although very often groups of apparently distinct species have virtually

|

46 ES Wie Cin O's Sika ¥,

identical terminalia. Throughout most of the Rutilza s.l. species the male sternite 5
does not significantly differ from that of other Rutilini, and has a simple bilobed
form (Text-figs 30 & 31), but in two aggregations of species (here treated as the
subgenera Grapholostylum and Microrutilia) the sternite is atypical: in Grapholo-
stylum the sternite has exceptionally pointed outer sides and has a pair of curious
blunt downwardly directed protuberances near the middle (Text-fig. 33); in
Microrutilia the sternite is also very prominent in profile when in situ but does not
have the submedian prominences and has each of the lateral lobes distinctly concave
on its hind margin (Text-fig. 32).

The aedeagus in Rutilia s.l. is remarkably constant in structure and the only
character found in it of any taxonomic use lies in the distiphallus. The distal
membranous part of the distiphallus is normally shorter than, or at most subequal
in length to, the sclerotized proximal part (Text-fig. 37), but in the subgenus
Grapholostylum the distal membranous part of the distiphallus is much longer than
in other Rutilia and has a slender whip-like form which is about twice as long as the
sclerotized proximal part of the distiphallus (Text-fig. 38). The form of the cerci
(mesolobes) does not differ very greatly in Rutilia s.1. species, although some specific
differences are evident, and the cerci do not provide features of subgeneric value.
On the other hand the shape of the surstyli (paralobes) differs conspicuously among
different subgenera, being particularly distinctive in the subgenera Donovanius and
Chrysorutilia. In Donovanius the surstyli are very enlarged and foliaceous in form
(Text-figs 66-71) (enabling males of this subgenus to be recognized immediately
the genitalia are examined), and in Chrysorutilia the surstyli (though varied in
shape) always come to a fine sharp point at the apex (Text-figs 72-84). Other
subgenera are less distinctive in genital form among tbeir included species, but they
all differ trom Donovanius and Chrysorutilia by having differently shaped surstyli
(i.e. by having neither the enormous foliaceous surstyli like Donovanius nor the
sharply pointed mucronate surstyli like Chrysorutilia).

In Rutilia s.l. as a whole it is often difficult, unless a species has a particularly
unusual form of male hypopygium, to determine the limits of species. Many of the
apparent species have geuitalia that are identical or nearly identical—for example,
the many distinctive-looking entities considered to be species in the subgenus
Donovanius actually differ not at all in their male genitalia, or if there are differences
they are extremely subtle and of no practical use. Some of the colour characters
that have been used to separate species, for instance whether the metallic areas of
the abdomen are in spots or bands, seem to be undependable criteria for they fail to
correlate very often with differences in male genitalia (particularly in the subgenus
Chrysorutilia in which some species with very distinctive genitalia show the same or a
very similar range of variation in colour pattern). Furthermore it seems possible
that some species are polymorphic in respect of hair colour, so that some apparent
species with yellow pleural hair may not be actually distinct from those with black
pleural hair; and it also appears likely that some species may have black pleural
hair in males but yellow pleural hair in females (similar to the sexual dimorphism in
pleural hair colour found in some species of the higher Tachinid genus Winthemia
Robineau-Desvoidy). At the present time no firm conclusions can be drawn on

REVISIONARY CLASSIFICATION OF RUTILIINI 47

these possibilities, so specific names are maintained as valid in the present work
unless the types on which they are based show complete agreement in their characters
(hair colour included). In taking this approach I have perhaps erred on the side of
accepting too many nominal species as valid, but in the present state of knowledge it
is better to do this than to establish synonyms on inadequate evidence of conspeci-
ficity.

KEY TO THE SUBGENERA OF RUTILIA

| Note: Occasional specimens may have four postalar setae on one side and three on the
other: such specimens should be run as if four setae were present on both sides. R. micropalpis
is subgenerically unplaced and will not therefore run out in the key (it is distinguished from
all other Rutilia s.1. by the very reduced palpi and other characters noted on p. 92).]

1 Parafrontals non-pollinose, brilliant metallic green to blue-violet. Postalar callus
with three setae. Hind tibia without anterodorsal fringe and without definite ad
or pd setae. [Eastern Australia] . : ; NEORUTILIA Malloch (p. 48)

— Parafrontals pollinose, either without metallic colour or with mere trace near vertex
(except in species from Philippines but then four postalar setae). Postalar callus
with either three or more setae. Hind tibia either with well developed close-set
anterodorsal fringe or with some well developed ad or pd setae, or with both fringe
and pd setae. : : :

2 Suprasquamal ridge bare ani postalar callus with four or five setae. Metallic green,
blue to purplish black forms with bold white pollen spots on thorax and abdomen.
{Eastern Australia] : , : AMENIAMIMA subgen. n. (p. 51)

— Suprasquamal ridge haired, or " bare then only three strong setae on postalar callus.

Colour and pattern varied, but if bold white pollinose spots present then either
three setae only on postalar callus or parafacials haired. 3

3 Postalar callus with four or more strong setae. Hind tibia w ith an antecedorsal
fringe. One sternopleural seta (o + 1), at most only a very weak anterior sterno-
pleural seta distinguishable among the hair. Hair of suprasquamal ridge long,
dense and bushy, often crinkled and often extending on to basal depression of
lower calypter 5 : . : : : ; ; : 4

- Postalar callus with three strong setae. Hind tibia without a definite anterodorsal
fringe (occasional species with weakly developed or irregular fringe). Two or three
sternopleural setae (I + 1 or 2+ 1), anterior st// conspicuous amongst the
sternopleural hair. Hair of suprasquamal ridge rather short and sparse, not
noticeably crinkled and never extending on to base of lower calypter (lower calypter
always entirely bare)

4 Last abdominal tergite (T5) with a inedian depression and a meee faeeverse row of
strong erect setae. ¢ genitalia with very large broad foliaceous surstyli (Text-figs
66-71) without sharp pointed tip. Scutellum usually distinctly flattened and
without distinct preapical setae in front of the marginal row. Pteropleural hairing
not developed in front of the level of the posterior stp/ seta (Text-fig. 20). Inner
half of humeral callus with at least a trace of humeral setae distinguishable from
the hair (humeral callus usually therefore with three or four distinguishable setae
even if the innermost one or two are very weak). Head with dark ground colour.
[Australia, Solomon Islands to Samoa] . : DONOVANIUS Enderlein (p. 69)

— Last abdominal tergite ([5) evenly convex across its width, without median depres-
sion or at most with only a mere trace of apicomedian hollowing; T5 normally
without any development of strong setae, vestiture composed only of fine erect
hair (some setae present in species from Philippines, and some other species occa-
sionally with some of the erect hair stronger than remainder). ¢ genitalia

tN

un

48 Re We CROSS KEY

with elongate surstyli which end in a sharp pointed tip (Text-figs 72-84). Scutellum
convex and with an irregular row of small but definite horizontal preapical setae
in front of the marginal setae. Pteropleuron haired on the anteroventral part in
front of the level of the posterior stp] seta (Text-fig. 19). Inner half of humeral
callus without setae developed amongst the hair (humeral callus therefore only
with the two setae on the outer half). Head usually with bright yellow ground
colour. [Widespread in Oriental and Australasian Regions]
CHRYSORUTILIA Townsend (p. 54)
5 Last abdominal tergite (T5) with a median depression. Abdominal T3 with a
transverse row of at least a few, usually many, strong erect spiniform marginal
setae. Scutellum distinctly flattened or slightly hollowed before the apex.
Suprasquamal ridge bare or haired. Arista micropubescent. Setae of inner
ventral ends of abdominal tergites rather strong and directed downwards as well as
backwards. <¢ sternite 5 with normal simple lobes. [Australia]
RUTILIA Robineau-Desvoidy s.str. (p. 77)
— Last abdominal tergite ([5) without a median depression, the upper surface evenly
convex or at most with only a trace of flattening at the tip. Abdominal T3 without
a transverse row of marginal setae, or if a row present then the setae not markedly
spiniform. Scutellum evenly convex on upper surface. Suprasquamal ridge
haired. Arista long-pubescent to short-plumose. Marginal setae of tergite
venters usually weak or hair-like and nearly completely recumbent (not projecting
noticeably downwards). <3 sternite 5 atypical, shaped either as in Text-fig. 32 or
as in Text-fig. 33 . 6
6 6 sternite 5 as in Text-fig. 3 By the eae wary Sirouely, Receaaneiee on each side ead
provided with a pair of submedian downwardly directed protuberances. Distal
membranous part of g aedeagus exceptionally long and whip-like (about twice as
long asthe sclerotized proximal part of distiphallus, Text-fig. 38). Only one
posterior intra-alar seta (exceptionally a small second seta present in front of main
one). Q normally without, or with one pair of, proclinate orbital setae. Thorax
with distinct white pollinose areas (often forming bold white spots) over meso-
pleuron, sternopleuron, humeral callus and supra-alar area of scutum. [Tasmania
to Queensland] . : : : GRAPHOLOSTYLUM Macquart (p. 83)
— dsternite 5 as in Text-fig. 32, the sternite without such acuminate sides and without a
pair of submedian protuberances. Distal membranous part of gj aedeagus normal
in size, shorter than proximal sclerotized part of distiphallus (Text-fig. 37).
Normally two post 1a setae (but only one in occasional specimens). @ normally
with two pairs of proclinate orbital setae (occasionally one or none). Thorax
without distinct white pollinosity and therefore lacking bold white spots. [Tas-
mania to Queensland; one undescribed species seen from New Guinea]
MICRORUTILIA Townsend (p. 86)

Subgenus NEORUTILIA Malloch

Neorutilia Malloch, 1936: 17. Type-species: Rutilia (Neorutilia) simplex Malloch, 1936, by
original designation. (As subgenus of Rutilia Robineau-Desvoidy, 1830).

Diacnosis. Parafrontals metallic (hardly at all pollinose). Epistome metallic; genal dila-
tions entirely metallic (trace of thin pollinosity in some lights). Facial carina flattened on outer
surface, strongly contracted (‘pinched-in’) ventrally in 9, not separated from lunula by distinct
depression. Parafacials bare. 2 without proclinate orbital setae. Arista bare (unusually
long and slender). Humeral callus with 4-5 setae. Posthumeral setae absent in g, variably
2-3 in 9 (one near inner edge of humeral callus and at least one near presutural seta). One
post ia seta in $, one or two small post ia setae in addition to main one in 9. Scutum with super-
numerary prescutellar setae, forming in all a transverse row of about 12 very strong stiff setae

eee

CE — ——— — ———————————————————
———————

REVISIONARY CLASSIFICATION OF RUTILIINI 49

immediately before scutellum. Postalar callus with 3 strong setae. Suprasquamal ridge
thickly haired. Scutellum strongly flattened, rather thin; with 7-8 pairs of marginal setae
(these very strong, stiff and straight); with a row of small irregular preapical setae, these very
feebly developed in §. Haired area of lower part of pteropleuron not extending in front of
level of posterior sternopleural seta. One sternopleural seta (o + 1). Prosternum and pro-
sternal membrane bare. Hind tibia without definite anterodorsal fringe and without ad setae,
pd seta absent in gf but one present in ? (latter may have very weak second fd setula). Last
abdominal tergite broad and with large depression. T3 without median or lateral marginal
setae. 15 with transverse median regular row of long strong erect setae. Surstyli of f genitalia
simple elongate lobes (Text-fig. 59). [Bright green to violaceous blue species with metallic para-
frontals and genae and blackish hind margins to abdominal tergites].

DISTRIBUTION. Known only from eastern Australia from Victoria to Queensland.

Discussion. Malloch erected Neorutilia as a subgenus of Rutilia for the single
aberrant species simplex Malloch. This is a curious and isolated species showing an
unusual combination of characters which prevent it from being assigned satisfac-
torily to any other subgenus, and I therefore agree with Malloch in placing simplex
separately from other Rutilia and am recognizing Neorutilia as a valid subgenus;
no other species fitting into the concept have been described.

The metallic blue or green parafrontals, epistome and genal dilations make
simplex a unique species amongst the whole Rutilia fauna of Australia, and it is
therefore very easily identified, but very similar forms superficially—having the
same type of largely metallic head, similar coppery green to blue-violet body colour,
and even the same knob-like development of the hind part of the notopleuron—are
to be found in the luzona-group of Chrysorutilia from the Philippines. This super-
ficial resemblance is so strong that I at first inclined to believe that simplex and the
Philippine species referred to must have strong affinity and perhaps be consubgeneric,
but detailed examination of the whole constellation of body characters shows con-
vincingly that the resemblances are convergent. The /uwzona-group has the features
of true Chrysorutilia such as the presence of strong hairing on the prosternal mem-
brane and forward edge of the prosternum itself, loss or non-development of the
inner setae of the humeral callus, no depression in the last abdominal tergite,
extended area of hairing forwards on the lower pteropleuron, no supernumerary
prescutellar setae, four or more postalar setae, convex scutellum, and long regular
hind tibial fringe: in simplex, however, these characters of Chrysorutilia are not
found, and Neorutilia differs from Chrysorutilia by having bare parafacials, bare
prosternal membrane and prosternum, inner humeral setae developed, strong
supernumerary prescutellars developed, only three postalar setae, extremely flattened
scutellum, pteropleural hairing not extending forwards of the sternopleural seta,
no hind tibial fringe (or extremely short and inconspicuous), and a well developed
depression in T5.

In several of the characters just cited Neorutilia resembles the subgenus Dono-
vanius—for example both subgenera have setae on the inner part of the humeral
callus, lack pteropleural hair anterior to the sternopleural seta, have a very broad
abdominal T5 with large median depression, and have the scutellum distinctly
flattened—but it is doubtful whether there is any close relationship between the two.
Neorutilia differs from Donovanius in having only three postalar setae, largely

50 R. W. CROSSKEY

metallic head, only one stfl seta and no hind tibial fringe; the hairing of the
suprasquamal ridge is also much longer, more bushy and crinkly, than in Donovanius
(in which respect Neorutilia agrees with Chrysorutilia). The hairing of the supra-
squamal ridge at once separates Neorutilia from Amenianuma subgen. n., from which
it also differs by the pteropleural character, the metallic head, the three postalars,
the lack of white pollinose spots, the flattened scutellum and the supernumerary
prescutellar setae.

In simplex the presence of very strong supernumerary setae on the scutum
immediately in front of the scutellum is a striking feature; it is difficult to distinguish
which actual setae are the supernumeraries and which are the true hindmost setae
of the dorsocentral and acrostichal rows, for together the setae (which are all un-
usually strong, rather straight and thick) form a transverse row of more or less
homogeneous setae (very much resembling then the transverse rows of prescutellar
setae formed in a similar way in many species of Formosia or in occasional species of
Rutilia (Donovanius) such as sabrata (Walker)). Development of such super-
numerary prescutellars appears to occur rather haphazardly in unrelated groups of
Rutilines.

It does not seem likely that Neorutilia has any close relationship with the other
subgenera possessing only three postalar setae, Rutilia s.str., Grapholostylum, and
Microrutilia, for apart from this character and the lack of the hind tibial fringe it
has little in common with these subgenera, differing from them greatly in general
appearance, metallic head, one stpl seta, flattened scutellum, supernumerary pre-
scutellar setae, and the long thick suprasquamal hairing (in contradistinction to the
rather short sparse ridge hairing found in Rutilia s.str. or Grapholostylum).

Apart from the characters cited in the diagnosis and discussed comparatively
above there are some other curious features found in simplex that require comment.
The facial carina seems, from the small number of specimens so far available, to be
unusually sexually dimorphic, being much longer and more strongly pinched-in
towards the lower end in the female than in the male, and the antennae of the female
to be correspondingly longer than those of the male. There is also something
‘different’ about the facies of the head when seen in facial view that is hard to
define in words or to figure satisfactorily but is associated with the fact that the
lower ends of the facial ridges are more flattened than is usual in Rutilia s.1. end more
extensively haired; normally in Rutilia there are only one or two irregular series of
hairs on the facials laterad and ventrad of the main vibrissae, but in simplex there
are some three or four irregular series of rather small hairs. Another unusual
feature is the nature of the arista, which is exceptionally long and fine and is virtually
totally bare; in all other Rutilia the arista is obviously more thickened on much of
its length and is always conspicuously pubescent or sometimes even short-plumose.
The claws are shorter than in most other species of the genus, and the fore tarsi are
shorter than in all other species. Malloch (1936) pointed out in the original definition
of Neorutilia and description of simplex that the section of the costa between the
apex of the subcosta and the first vein (second costal sector) is subequal in length
to the section between the apex of the first vein and apex of the second vein (third
costal sector), and remarked that he had not seen other species with similar costal

REVISIONARY CLASSIFICATION OF RUTILIINI 51

divisions. Although I have not used this feature in the subgeneric diagnoses, it is
true of all material that I have seen that the second and third costal sectors in
simplex are subequal in length whereas in all other Rwtzlza the second sector is ob-
viously appreciably shorter and often very much shorter than the third sector;
certainly this is one of the characters along with the others already alluded to that
make simplex a very atypical Rutilia, and one best placed in a separate subgenus.
INCLUDED SPECIES

Rutilia (Neorutilia) simplex Malloch. AusTRALIA (New South Wales, Queens-
land, Victoria). [Holotype examined].

Subgenus AMENIAMIMA subéen. n.
Type-species: Rutilia argentifera Bigot, 1874.

Diacnosis. Parafrontals thickly pollinose. Epistome not metallic, genal dilations thickly
pollinose. Facial carina flattened on anterior surface, sides subparallel or slightly widened near
base of third antennal segment, not separated from lunula by distinct depression; epistome
rather strongly prominent. Parafacials bare. 9 without proclinate orbital setae or with one
pair. Arista pubescent. Humeral callus with 3-4 setae, innermost one well developed
(especially in 9). Posthumeral setae usually absent in $ or one distinguishable from hair near
inner edge of humeral callus, 2-4 rather strong but irregular in @ (if more than two then one of
the ph standing near to presutural seta). One or two post ia setae. Scutum without super-
numerary prescutellar setae. Postalar callus with 4-5 strong setae. Suprasquamal ridge
bare. Scutellum not flattened, dorsal surface evenly convex; with 3-6 pairs of marginal setae
(these not stiffened) ; marginal setae preceded by irregular row of distinct preapical setae (very
well differentiated from scutellar hairing). Haired area of lower part of pteropleuron extending
well forwards of the posterior stpl seta. Normally two sternopleural setae (1 + 1), anterior
one weak in f and sometimes not differentiated in this sex (aberrant 2? specimen seen with doubled
posterior stpl). Prosternum and prosternal membrane bare. Hind tibia without definite
anterodorsal fringe and with one submedian ad seta (except in cingulata § which has moderately
developed regular ad fringe), with one or two fd setae. Last abdominal tergite (T5) with a
very small median apical depression in $ (not longer than half tergite length at most) and
without any definite apical depression in 9. T3 without median marginal setae (occasionally
a few developed in 2) and with lateral marginal setae. T5 with a transverse row of long strong
erect setae on apical half, sometimes whole posterior half of T5 in $ bristled with many very
long strong irregularly arranged setae. Surstyli of $ genitalia in form of broad lobes (Text-fig.
58). [Bright green, cupreous, or blue to purplish black forms with bright orange-yellow heads,
and discrete bold white pollinose spots on thorax and abdomen, showing strong resemblance to
calliphorid flies of genus Amenia Robineau-Desvoidy].

DistTRIBUTION. Australia only, from Victoria to Queensland (not seen from
Western Australia, South Australia, or Tasmania).

Discussion. The new subgenus Ameniamima is here erected for a small group
of eastern Australian species of Rutzlia s.1. which show a quite remarkable convergent
resemblance in body form, colour and pattern, to flies of the Australian genus
Amenia Robineau-Desvoidy (Calliphoridae). Four nominal species of this group
have been described, but each is known from only very little material and it is not
certain that all four names actually denote distinct species; one is here placed as a
new synonym.

52 R. W. CROSSKEY

Ameniamima includes all the species of Rutilia s.l. in which the suprasquamal
ridge is bare and there are more than three postalar setae (elsewhere in Rutilia s.l.
the few species with bare ridge have only three postalars). The metallic green or
blue colouring that is typical in Ameniamima, combined with the bright orange-
yellow head, give the species a superficial resemblance to some Chrysorutilia species,
and it is possible that it is to this latter subgenus that Ameniamima is most closely
related; the two subgenera share such features as the presence of four or five postalar
setae, the haired area of the lower pteropleuron extending well forwards of the
level of the posterior sternopleural seta, the presence of a definite irregular transverse
row of semi-horizontal preapical scutellar setae in front of the marginals, similar
general shape of facial carina, and similar lack of a definite median depression in the
last abdominal tergite (though a small depression is present at the tip of the tergite
in Ameniamima males). The main differences between Ameniamima and Chryso-
vutilia, apart from the bare suprasquamal ridge in the former and densely haired
ridge in the latter, include the following: parafacials, prosternum and prosternal
membrane bare in Ameniamima (partly or entirely haired in Chrysorutilia) ; inner
half of humeral callus with one or two humeral setae developed in addition to the
pair on the outer half of the callus in Ameniamima (only the outer pair present in
Chrysorutilia) ; hind tibia usually without definite fringe in Ameniamima (with long
close-set regular fringe in Chrysorutilia); abdominal T5 with long strong setae in
Ameniamima (only with fine hairing in Chrysorutilia, except for some weak erect
bristling in Philippine forms). In addition the chaetotaxy is generally much stronger
in Ameniamima than in Chrysorutilia, especially the bristling of the mesonotum,
the outermost humeral setae, and the posterodorsal setae of the hind tibia. The
head of Ameniamima species is almost entirely thickly yellow pollinose over the
yellow ground colour, only the extreme upper parts of the occiput being bare or |
metallic, and the completely pollinose genal dilations and postbuccae will at once |
distinguish specimens of the subgenus from Chrysorutilia (in which all species have
these areas of the head largely bare or metallic and at most only thinly pollinose).

The body colour in Amenianuma ranges from brilliant metallic green (sometimes
with brassy or coppery tinge) to blue, violet and purplish black; in one species,
cingulata, the colour is virtually black except for violet tinges near the pollinose spots
and metallic silvery blue colour underlying a thick whitish pollinosity on abdominal
T3. All the species possess well developed, conspicuous and rather discrete, spots
of thick white pollinosity, those of the abdomen having a slightly shifting appearance
with the light. The spot pattern consists of paired white spots in these positions:
humeral; supra-alar; mesopleural; sternopleural; ventrolateral on abdominal T3,
T4, and T5; dorsolateral on T3; submedian on T4 and T5. In addition there is
usually a pair of submedian white pollinose vittae variously developed on the
prescutum, and in a few specimens (including the lectotype of argentifera) there is a
trace of an evanescent pair of submedian spots on T3. Usually the dorsolateral
spots of T3 and the submedian spots of T5 are more or less merged with the ventro-
lateral spots of their respective tergites. In the male of cingulata, which conforms
to the basic plan of the pattern, the supra-alar spots and the submedian spots of
T4 and T5 are rather small but the white pollinosity is so extensively developed on

> |

REVISIONARY CLASSIFICATION OF RUTILIINI 53

the ventrolateral and dorsal parts of T3 that it forms an almost complete band around
the whole of this tergite and in dorsal view completely conceals the metallic blue
ground colour of this tergite.

A white-spotted pattern of the type just described does not occur in quite the same
way in other subgenera of Rutilia s.1., although some of these contain individual
species in which there is some development of a white-spot pattern with a superficial
resemblance to that of Ameniamima, for example an undescribed species of subgenus
Chrysorutila in which the thoracic spotting is the same except for lack of definite
sternopleural spots and in which the abdomen has ten small sharply-defined silvery
white spots on a purplish black background (four spots in a transverse row on each
intermediate tergite, and paired lunate spots on the last tergite); two species of
Grapholostylum have a pattern of whitish spots, viz. albovirida Malloch and
| dorsomaculata (Macquart), but in these species there are ventral spots or pollinose
areas on the tergites and a pair of submedian spots on the scutum which are not
found in Ameniamima. The resemblance, however, between the spot-pattern of
Ameniamima and of the Calliphorid fly genus Amenia is extraordinarily exact, and
the former name here proposed for a subgenus of Rutilia alludes to this fact as well
as to the other many similarities by which Ameniamima counterfeits Amenia.

The astonishing convergent resemblance between Rutiliini and Ameniinae
(Calliphoridae) was already recognized in an inchoate way as early as 1830, when
| (as Crosskey (1965 : 41) has pointed out) Robineau-Desvoidy realised that it was by
an error that he had placed the Ameniine species Amenia leonina (Fabricius) in
Rutilia when the latter was first described. Since then several workers have com-
mented on the resemblance and some have been misled into uniting the Ameniines
with the Rutiliines, mainly because of the common possession of a large facial carina
_and the same range of colour and pattern; Enderlein (1936) named the Ameniine
| genus I’ormositomima for a species seemingly mimicking the black-and-white pattern
of Formosia speciosa (Erichson). It is now established beyond any doubt that the
_ Ameniines, with their Calliphorid type of male genitalia (completely unlike the
, Prosenine Tachinid genitalia with the extraordinarily elongate slender aedeagus) and
| other Calliphorid characteristics are not Tachinidae (see Crosskey (1965)), and hence
' that the resemblances are due to convergence. These resemblances reach their
/ apogee amongst the species of Rutilia here placed in the subgenus Ameniamima

and in the species of Amenia Robineau-Desvoidy, in which it almost seems as though

individual species have their respective counterparts: bright green specimens of
| R. (A.) argentifera have their counterpart in Amenia imperialis R.-D., while R. (A.)
| quadripunctata (Malloch), which is often more blue, resembles A menia leonina (Fab.).
| However, all the Amenia species have a pair of silvery white pollinose spots in a
| postalar position (partly overlying the postalar callus and partly the hind scutal
| border) which are always absent in Rutilia s.l.
_ Three of the nominal species included in the present subgenus were placed by
Malloch (1929, 1930) when originally described in the genus Formosia because of the
bare suprasquamal ridge, a character which Malloch rather rigidly regarded as
‘absolutely dependable for separating Formosia from Rutilia; reliance on this charac-
| ter alone induced Malloch to place the three ‘bare ridge’ species here included, and

54 R. W. CROSSKEY

also confusa Malloch (a species of Rutilia s.str. with bare suprasquamal ridge), out of
Rutilia to which they so obviously fit well on the total facies provided by all their
other characters and into Formosia (with which, apart from the bare ridge, they
have little in common). The oversimplified picture provided by Malloch’s concep-
tion of two genera Formosia and Rutilia always clearly separable by the ridge
vestiture character is not justified when all characters are correlated, and the species
here forming the subgenus Ameniamima are considered in this revision to belong to
Rutilia s.l. in spite of the exceptional feature of lacking hair on the suprasquamal
ridge.

Note should be made here of Malloch’s (1928a : 333; 1929 : 297; 1930 : 107)
misidentification of Rutilia argentifera Bigot. The true argentifera Bigot with bare
prosternal membrane, bare parafacials and bare suprasquamal ridge, is the species
here designated as type-species of Ameniamima subgen. n. and is not the species called
argentifera by Malloch (who did not see Bigot’s types and whose citations given
above to this species are all based on misidentifications). Malloch’s species to which
he applied Bigot’s name has the prosternum hairy (see Malloch’s 1928a : 333 key)
and the parafacials haired (Malloch 1929 : 297-298), and also has the suprasquamal
ridge hairy (otherwise he would have placed it in Formosia), and is clearly a species
of the subgenus Chrysorutilia.

INCLUDED SPECIES

Rutilia (Ameniamima) argentifera Bigot. AUSTRALIA (New South Wales,
Queensland). [Lectotype examined].
Jrontosa (Malloch) syn. n. [Holotype examined].
R. (A.) cingulata (Malloch). AustrRALt1A (New South Wales). [Holotype
examined].
R. (A.) quadripunctata (Malloch) comb.n. AUSTRALIA (Queensland, Victoria).
[Holotype examined].

KEY TO SPECIES OF THE SUBGENUS AMENIAMIMA

1 Postorbits entirely bright yellow pollinose and concolorous with rest of the yellow

head. ¢ surstyli as in Text-fig. 58 . - . R. argentifera
— Upper two-thirds of the postorbits silvery white pollinose and contrasting in colour
with rest of the yellow head. [Genitalia of Jf not examined] : 2

2 Third abdominal tergite (i.e. apparent second segment) almost completely coven!
with dense white pollinosity, so that abdomen shows a broad pale transverse sub-
basal band which strikingly contrasts with remainder of dark abdomen (silvery
bluish glints visible through the pollinosity in some lights). General body colour
dark purplish black with very dark brown non-metallic scutellum. [Known only
from g holotype, 9 may not conform] - : : R. cingulata
- Third abdominal tergite not so, with only two small niger spots of whitish
pollinosity with shifting appearance on each side, T3 therefore appearing unicolorous
with rest of abdomen. General body colour blue-green to deep violet-blue,
scutellum distinctly metallic . : c : : : : R. quadripunctata

Subgenus CHRYSORUTILIA Townsend stat. n.

Chrysorutilia Townsend, 1915 : 23. Type-species: Rutilia formosa Robineau-Desvoidy, 1830,
by original designation.

REVISIONARY CLASSIFICATION OF RUTILIINI 55

Philippoformosia Townsend, 1927 : 282. Type-species: Philippoformosia splendida Townsend,
1927 [=townsendi Crosskey, nom. n.], by original designation. Syn. n.

Habrota Enderlein, 1936 : 399. Type-species: Rutilia foymosa Robineau-Desvoidy, 1830, by
original designation. [Isogenotypic name with Chrysorutilia Townsend].

Zovamsceus Enderlein, 1936: 416. Type-species: Rutilia erichsonit Engel, 1925 [=Dexia
chersipho Walker, 1849], by original designation. Syn. n.

Idania Enderlein, 1936 : 408. Type-species: Idania atrox Enderlein, 1936, by original designa-
tion. Syn. n.

Formotilia Paramonov, 1968 : 355. [Name published in footnote to generic key: unavailable
under Article 13 (b) of International Code of Zoological Nomenclature, no fixation of type-
species. ]

Diacnosis. Parafrontals pollinose (except /uzona-group and atvox). Epistome not metallic
(except Juzona-group). Genae with the dilations partly or entirely metallic, without or with
very thin pollinosity (golden green to blue, except dark brown in atyov). Facial carina
conspicuously flattened on anterior surface, sides subparallel or slightly divergent ventrally,
separated from lunula by weak depression. Parafacials haired. 9 without proclinate orbital
setae. Arista micropubescent. Humeral setae reduced, 2 on outer half of callus only (at most
only minute hair-like inner humerals developed). Posthumeral setae absent. Post ia setae
normally one each side, occasionally absent (rarely two in 9). Scutum without supernumerary
prescutellar setae. Postalar callus with 4-6 strong setae. Suprasquamal ridge thickly haired.
Scutellum not flattened or at most with only slight flattening at apex; with 4-7 pairs of marginal
setae (these rarely stiffened); marginal setae preceded by an irregular row of horizontal pre-
apical setae clearly differentiated from scutellar hair. Haired area of lower part of pteropleuron
extending well forwards of the posterior stp] seta. One sternopleural seta (o + 1), sometimes
one or two small wiry anterior sternopleurals developed in 9; rarely sternopleurals quite absent.
Prosternum haired on anterolateral corners; prosternal membrane haired (hair often long and
conspicuous). Hind tibia with long regular anterodorsal fringe (no evident ad setae), without
or with one or two very weak fd setae. Last abdominal tergite ([T5) without median depression,
convex on upper surface. T3 without median marginal setae (rarely weak setae present in 9)
and without lateral marginals. T5 with fine erect hair only (sometimes some more strongly
developed slightly setiform vestiture at apex; irregular erect stubby setae in Juzonag). Sternite
5 of g with simple rounded lobes. ¢ genitalia with distal membranous part of distiphallus
shorter than, or at most subequal in length to, the sclerotized proximal part; surstyli much longer
than wider, sometimes with blunt projection or long tooth anteromedially, always with sharp
pointed tip that is directed more or less forwards (Text-figs 72-84). [Mainly metallic golden-
green, coppery, or blue-green forms with banded or spotted abdomen and yellow head ground
colour, often with white pollinose areas on thorax but these very rarely forming bold discrete
spots. ]

DIsTRIBUTION. Widespread throughout Oriental and Australasian regions, the
distribution including Ceylon, India, Vietnam, Philippines, Timor, Western
Australia (including Monte Bello Islands), all of eastern Australia and Tasmania, the
Kei Islands, New Hebrides and Lord Howe Island.

Discussion. Chrysorutilia is the largest subgenus of Rutilia s.1. and has a wider
geographical distribution than any other subgenus; it includes some two-fifths of the
described species (though some of the names accepted as valid might prove to be
synonyms) and is the only segregate of Rutilia s.1. represented on the mainland of
Asia. The subgenus is poor in number of species in the Oriental Region, and those
that do exist in this area appear to be uncommon to judge from the paucity of
specimens in museum collections; the occurrence of the subgenus in Asia appears
to be due to dispersal from a centre of evolution in Australia, and the forms found

56 R. W. CROSSKEY

in the New Hebrides and Lord Howe Island (whose specific identities are doubtful
at present) probably also reached these locales from Australia itself.

The subgenus is perhaps better characterized than the other subgenera and there
is usually no difficulty in deciding whether any described species does or does not
fit into Chrysorutilia. The subgenus includes all the Rutilia species that have bright
yellow heads and hairy parafacials, and it is a useful rule-of-thumb that specimens
showing these characteristics in combination must belong to species of Chrysorutilia;
however, there are a few species which evidently belong in this subgenus in which
the head is not bright yellow (even formosa, the type-species, has the head colour
slightly brownish rather than yellow) and a few very curious forms from the Philip-
pines in which the body is entirely blackish brown (atrox Enderlein) or in which the
parafrontals are bare and brilliant metallic green or blue (Juzona Enderlein and
splendida Townsend). These Philippine species are discussed further below, as
two of them are type-species of generic names which are here treated as synonyms of
Chrysorutilia.

The characteristics which separate Chrysorutilia from Donovanius and Amenta-
mima are discussed in the treatments of these subgenera and need not be detailed
here, but some unusual features of Chrysorutilia are worthy of special note. A
curious character of the subgenus is the unusually long and conspicuous hair to be
found on the membranous areas of the prosternal region, and anteriorly on the edge
of the prosternal plate itself; hair on the membrane occurs in many Donovanius
species and occasionally in Grapholostylum, but in these subgenera is usually sparse
and not very obvious, while they never have hair actually on the edge of the proster-
num, whereas in Chrysorutilia both membrane and plate bear hair and that on the
membrane is sometimes strong and black and immediately obvious (providing the
head of the specimen is set forward sufficiently clear of the lower thorax). Prosternal
membrane hair of this strong black type occurs on the types of Idania atrox (type of
Idania) and Philippoformosia splendida Townsend (type of Philippoformosia), and
conformity of this character (as well as other essential features) suggests that these
Philippine species should be treated as consubgeneric with typical Chrysorutilia;
hence Idania and Philippoformosia are here placed as new synonyms of Chryso- ~
rutilia.

Idama and Philippoformosia agree, too, with Chrysorutilia in other essential
features of the subgenus such as the rounded non-sulcate last abdominal tergite,
the haired area of the pteropleuron extending well forwards of the posterior sterno-
pleural seta, the weak development of humeral setae (only two setae on the outer
half of the callus), the total lack of posthumeral setae, the extremely well developed
hind tibial fringe, the very bushy tuft of rather long crinkly hair on the suprasquamal
ridge, and the presence of distinct preapical setae on the scutellum just in front of the
marginals. However, there are some obvious minor differences between the
Philippine species and typical Chrysorutilia and they are here placed in separate
species-groups from the main bulk of the species; the groups are keyed out and
defined immediately after this discussion section.

Hairing of the suprasquamal ridge is developed to its greatest extent among the
Rutiliini in this subgenus. In all Chrysorutilia species the hair of the ridge is very

REVISIONARY CLASSIFICATION OF RUTILIINI 57

long and dense and forms a very conspicuous bushy tuft in which the individual
hairs are often rather crinkled apically (Text-fig. 24) ; in some of the species the hairi-
ness extends on to the base of the lower calypter. Hairing of the lower calypter
occurs, so far as I know, nowhere else in the Rutiliini, but as it is not at all consistent
throughout Chrysorutilia it has not been cited as a character either in the subgeneric
key or the diagnosis; nevertheless it can be a useful feature in recognizing, or helping
to distinguish, some species. In the smaller species of Chrysorutilia the lower
calypter is usually completely bare or has only a very few hairs at the extreme base
adjacent to the suprasquamal ridge, but in some of the larger species such as R. (C.)
splendida (Donovan) the whole basal depression of the calypter has conspicuous erect
dark hair on the upper surface; in R. (C.) cryptica sp. n. half the surface of the lower
calypter or more is hairy.

In Chrysorutilia the genal dilations of the head are much more extensively metallic,
because they are only very thinly pollinose or because they are bare, than in the
other subgenera; in other Rutilia s.1. the lower parts of the head are thickly pollinose
(except in Neorutilia) and only the postbuccal regions at most have any trace of
metallic colouring. In Chrysorutilia at least the posterior half and sometimes the
whole of the genal dilation is shining golden green, cupreous, blue-green or blue-
violet, at least in some lights. In the Jwzona-group even the epistome is largely
shining golden green to steely blue and only very thinly pollinose, and there are bare
shining parafrontals. MHairing of the parafacials is more developed than in other
Rutilia s.1., and the males of many Chrysorutilia species have specially dense hair
which reaches to the bottom end of the parafacials, to a point about level with the
bottom-most point of the eye seen in profile; when hairy parafacials occur in other
subgenera the hairing normally does not reach so far down on the parafacials. The
facial carina in Rwutilia s.1. reaches its most wide and flattened condition in Chryso-
vutilia, with the result that the antennae in this subgenus tend to be distinctly more
widely separated at their insertions than is normally the case in other forms; nor-
mally the carina is exceptionally parallel-sided, tending not to widen at a level with
the base of the third antennal segment as in other subgenera, and often it even
widens noticeably towards the ventral end.

Three species-groups are recognized in Chrysorutilia and can be distinguished by
the following key:

KEY TO THE SPECIES-GROUPS OF CHRYSORUTILIA

1 Body extensively metallic, gold-green or cupreous to blue-violet . : 2
— Body uniformly blackish brown, not metallic, at most only slight trace of Paria
tinge on abdomen. [Philippines] . : atrox-group (p. 58)

2 Parafrontals and epistome metallic. Two posterior notopleural setae on unusually
strongly developed (almost knob-like) protuberance. [Philippines]
luzona-group (p. 58)
— Parafrontals and epistome not metallic. Notopleuron normal, with one posterior
seta and hind part of notopleuron sag prety et gayle {Not known from
Philippines] . : : : 2 formosa-group (p. 59)

58 R. W. CROSSKE Y¥

THE ATROX-GRouP

Diacnosis. Body blackish brown with at most only very faintest metallic purplish tinge
on abdominal tergites. Parafrontals non-pollinose, but not metallic. ¢g claws rather short and
thick. [Character of notopleural setae uncertain, one posterior seta on one side and two on
other in atyvox holotype, only specimen known.] Last abdominal tergite with preapical trans-
verse row of fine long erect setae differentiated from the hair.

This group contains only one species which Enderlein (1936) made the type-
species of his genus Idania. Though this species, atvox, is strikingly different from
all other Chrysorutilia because of the unicolorous dark mahogany-brown to blackish
brown colouring it agrees in all its essential features with the more typical species of
Chrysorutilia and I unhesitatingly place it in this subgenus ([danza thus sinking as a
synonym). At present, only the holotype of atrox is known; in some lights the
specimen shows very faint traces of purple on the abdomen, and the genae and
parafrontals—although brown and non-metallic—are seen to be devoid of definite
pollinosity (atvox in this respect resembling /uzona-group). The holotype shows
extremely well developed long black hair on the prosternal membrane and some
shorter fine hairs on the front of the prosternum, much as in Juzona-group, and the
hind tibial fringe is extremely well developed; it is a large specimen measuring some
20 mm in length with slightly infuscate wings in which the veins of the middle region,
especially anteriorly, are conspicuously yellow. It is possible that intermediate
forms will ultimately be discovered which interconnect atrox with the /uzona-group,
in which case the two groups could be merged under the latter name; at present both
groups are found only in the Philippines, an area from which typical Chrysorutilha
species (formosa-group) are not yet known.

INCLUDED SPECIES

Rutilia (Chrysorutilia) atrox (Enderlein) comb. n. Puitippines. [Holotype
examined].

THE LUZON A-GROUP

Diacnosis. Body largely golden green to dark blue; parafrontals, epistome and genal
dilations metallic. ¢ claws rather short and thick. Notopleuron with posterior part extra-
ordinarily protuberant and with two setae (notopleuron therefore with total of three setae).
Last abdominal tergite with transverse preapical row of long setae differentiated from the hair.

It is to this group that Townsend (1927) applied the name Philippoformosia, a
genus based on sflendida Townsend from the Philippines. As with atvox (above),
it seems to be that splendida Townsend has all the main characters shown by
Chrysorutilia and I therefore place the generic name based upon it as a synonym;
however, splendida together with /uzona Enderlein differs from typical Chrysorutila
by the features mentioned in the diagnosis above and separate species-group status
within Chrysorutilia is considered to be the most appropriate taxonomic treatment.
Unfortunately splendida Townsend is known only from the female holotype and
luzona only from the male holotype, and it is therefore difficult to be certain whether
two distinct species actually occur; Juzona should perhaps be placed as a synonym
of splendida Townsend, but for the present it appears best to maintain the names for

REVISIONARY CLASSIFICATION OF RUTILIINI 59

valid species pending more evidence. The name splendida Townsend, being herein
applied to a species of Rutilia s.l., becomes a junior secondary homonym of
R. splendida (Donovan), and the name townsend1 is therefore here proposed for the pre-
occupied splendida Townsend (the new name will enter synonymy with Juzona
Enderlein if it is later shown that the types of splendida Townsend and luzona
Enderlein are conspecific).

The BMNH collection contains two females of this group in rather bad condition
that were formerly in Bigot’s collection and are from an unknown locality (though
Philippines seems probable). These females were the specimens for which the late
Dr Paramonov had intended formally to erect the genus Formotilia, and which were
| the basis for the entry ‘gen.nov. No. 2’ and the footnote giving the published name
| Formotilia in his posthumous paper (Paramonov, 1968) ; one of the specimens bears a
| name label in Paramonov’s writing on which the generic name Formotilia is given.
This name, though published, is unavailable in nomenclature (no fixation of a type-
_ species), but the existence of the specimens in BMNH which were seen and labelled
| by Paramonov enables the name Formotilia to be placed. It clearly applies to the
| luzona-group here defined and if ever validated nomenclaturally would be a synonym
of Philippoformosia.

INCLUDED SPECIES
Rutilia (Chrysorutilia) luzona (Enderlein) comb. n. PutLippINes. [Holo-
type examined].
R. (C.) townsendi nom. n. PHILIPPINES. [Holotype of splendida examined].
splendida (Townsend) [Junior secondary homonym in Rutilia].

Tue FORMOSA-GrRoupP

Diacnosis. Body partly or largely metallic, golden green to blue violet; parafrontals and

_ epistome not metallic, genal dilations partly or largely but not completely metallic. ¢ claws

long and slender. Notopleuron normal, posterior part not exceptionally protuberant and with

one seta. Last abdominal tergite with fine erect hairing only, hair scarcely ever differentiated
into any definite erect setae.

_ This group contains all the Chrysorutilia species except for those few forms from
the Philippine Islands already discussed above; the formosa-group seems to be
absent from the Philippines and replaced there by the atvox and luzona groups, in
which there has been more extensive development of the bare or metallic areas of the
head, reduction of the male claws, and the development of some definite setae among
the hair of the last abdominal tergite (and also usually a doubling of the posterior
notopleural seta together with some exceptional swelling of the hind part of the
_notopleuron itself). In the formosa-group there are always only the normal two
notopleural setae (anterior and posterior), the parafrontals are thickly pollinose,
and the genal dilations (though partly metallic) are dull and thinly pollinose at least
anteriorly.

The concept of this group and of the subgenus Chrysorutilia as a whole rests
‘nomenclaturally on the identity of Rutilia formosa Robineau-Desvoidy, of which
| the original material is lost. When Townsend (1915) erected the genus Chrysorutilia

60 R. W. GCROSSEKE Yi

he cited no characters for it, but based it upon the nominal species R. formosa R.-D.,
for which a neotype is herein designated. As Townsend (1915) cited no characters
of formosa it is herein presumed that no conflict exists between formosa R.-D. as
here fixed by neotype (which equals formosa in Malloch’s and Enderlein’s sense)
and Townsend’s oviginal meaning of this name. For further discussion of this see
under neotype designation for R. formosa on p. 124.

The limits of species in this group are often very uncertain. Some apparently
distinct species which possess external pattern differences or differences of hair colour
have male genitalia that differ but slightly or not at all. On the other hand, some
very distinctive shapes occur in the male genitalia, especially in the shape of the
surstyli, and where unusual surstylus shapes occur they appear to be constant and to
define discrete species (for example, the bizarre shape of the surstylus in R. cryptica
sp.n. is unmistakable and constant). In the past species have usually been founded
on observed differences in external colour and pattern without regard to genital
characters, and names have been bestowed on supposed ‘species’ in which the
abdomen shows continuous transverse bands across the tergites and other names
given to ‘species’ in which the tergite pattern is formed of completely or partially
isolated spots. It has been found in the course of the present work that in many of
the Chrysorutilia species there is little or no correlation between the male genital
characters and the presence or absence or banded or spotted patterns. By removing
the genitalia of a large number of male specimens of R. (C.) spblendida (as this species
was previously understood) it has been found that three species have been confused
under this name which are easily recognized by constantly different shapes of the
male surstyli: in the commonest one (for which the name sflendida is fixed by
neotype) the surstylus is simple in lateral and posterior view, having no trace
of an anteromedian process (Text-fig. 72); in the second species (for which the name
decora is fixed by neotype) the surstylus has a characteristic blunt swelling antero-
medially which is just visible in posterior view (Text-fig. 73) ; and in the third species
(which in the absence of an available name is here newly described as cryptica) the
surstylus is produced anteromedially into an enormous forwardly directed tooth
and is strongly excavate and acuminate between this tooth and its apex (Text-fig. 74)
(the large tooth is very conspicuous in posterior view also). There is only a weak
correlation in these three species between the genital form and the spot-pattern or
band-pattern of the abdomen: in splendida the abdomen is usually banded but may
have the bands broken into discrete spots, in decora the abdomen usually has discrete
spots but may occasionally have continuous bands on one or both of the intermediate
tergites, and in cryptica the pattern is similar to decora (most often spotted but banded
specimens occurring).

Another example of more than one species confused under a single name is that of
R. (C.) imperialis. When males of this ‘species’ were examined for the present work
it was found that their genitalia had two distinct forms: in one the surstylus has a
blunt anteromedian projection (similar to that of decora) and the cerci are rather
slender in profile with the tips bent slightly forwards (Text-fig. 75); in the other the
surstylus is much narrower and has no anteromedian projection, and the cerci in
profile are broad medially with a rather sudden contraction before the rather straight

REVISIONARY CLASSIFICATION OF RUTILIINI 61

apical part (Text-fig. 76). These differences appear to be constant, and to define
two semi-sibling species, which cannot be differentiated on their external charac-
teristics. The original type-material of imperialis is lost and a neotype herein
designated fixes the name to the species with the genital form shown in Text-fig. 75.

The existence of species which can only reliably be separated by male genitalia
makes the determination of females rather hazardous, and it is not always possible
to make completely dependable associations of males with females. As some of the
primary types of described species are males and others females it is possible that
more of the names are synonyms than are given in the list of included species which
follows. Furthermore it is not yet clear whether pleural hair colour can be sexually
dimorphic in any species, or whether hair colour can be variously black or yellow
in the same sex in any species. At present black-haired and yellow-haired forms are
assumed to appertain to different species, but this assumption may not be substan-
' tiated in future when better criteria can be found for determining what is a species
than we have at present (at least it appears, so far, that a distinctive form of male
genitalia is correlated with a particular hair colour).

INCLUDED SPECIES

Rutilia (Chrysorutilia) caeruleata (Enderlein) comb. n. AvuSTRALIA (Western
Australia). [Lectotype examined. ]}
lineata (Enderlein) syn. n. [Lectotype examined. ]
R. (C.) caesia (Enderlein) comb. n. AvustTrRa.ta (Northern Territory, Queens-
land, Western Australia). [Lectotype examined. ]
vufibarbis (Enderlein) syn. n. [Lectotype examined.]}
viridescens (Enderlein). [Holotype examined. ]
R. (C.) chersipho (Walker) comb. n. AusTRALIA (Western Australia). [Neo-
type examined. |
evichsoni Engel syn. n. [Lectotype examined. ]
R. (C.) corona Curran. AvusTRALIA (New South Wales). [Holotype examined.]
R. (C.) cryptica Crosskey sp.n. AUSTRALIA (New South Wales, South Australia,
Victoria). [Holotype examined. ]
R. (C.) decora Guérin-Méneville. AusTRALIA (Tasmania to Queensland). [Neo-
type examined. |
R. (C.) formosa Robineau-Desvoidy. AusTRALIA (A.C.T., New South Wales,
Victoria). [Neotype examined. ]
pubicols Thomson syn. n. [Lectotype examined. ]
subvittata Malloch.
uzita (Walker) syn. n. [Holotype examined.]
R. (C.) goerlingiana (Enderlein) comb. n. AvusTRALIA (Western Australia).
[Lectotype examined. ]
R. (C.) idesa (Walker). AusTRALIA (? state). [Holotype examined.]
R. (C.) imperialis Guérin-Méneville. AustTRALIA (Tasmania to Queensland).
[Neotype examined. ]
ruficornis Bigot syn. n. [Holotype examined.]
semifulva Bigot syn. n. [Lectotype examined.]

62 R. W. CROSSKEY

R. (C.) imperialoides Crosskey sp. n. AUSTRALIA (Victoria to Queensland).
[Holotype examined].
R. (C.) nana (Enderlein)comb.n. Kat{[=Ker|Istanps. [Holotype examined. ]
R. (C.) panthea (Walker). AusTRALia (South Australia, Western Australia).
[Holotype examined].
R. (C.) rubriceps Macquart. AUSTRALIA (Queensland, ? Tasmania), CEYLON,
Inp1A, VIETNAM, Buru, ? Timor. [Holotype examined].
angustigena (Enderlein) syn. n. [Lectotype examined].
serena (Walker) syn. n. [Neotype examined].
nitens Macquart. [Holotype examined].
formosina Curran, 1930 syn. n. [Holotype examined].
R. (C.) splendida (Donovan). AUSTRALIA (Victoria to Queensland). [Neotype
examined].
australasia Gray. [Type-material lost: synonymy established by Walker
(1849 : 863) considered correct].
confluens (Enderlein). [Lectotype examined].
evanescens (Enderlein). [Holotype examined].
R. (C.) transversa Malloch. AusTRALIA (Western Australia). [Holotype
examined].

KEY TO SPECIES OF THE SUBGENUS CHRYSORUTILIA

[Note : The following key excludes the three species from the Philippines which are placed in
separate species-groups and can be recognized from the group characteristics given elsewhere.
The key attempts to place only the described species which are recognized in the present
revisionary classification: specimens of undetermined, possibly new, species exist in museum
collections and will not necessarily run out in the key (apart from a few species with distinctive
male genitalia, such as cryptica sp. n., there is still much doubt about specific limits in this
subgenus, and the supposed species tend to intergrade).]

I Hair of pleural regions of thorax pale yellow to golden orange. Hair of suprasquamal
ridge yellow. Hair of fore coxae partly or entirely yellow to golden red.
Mesonotum of ¢ with much pale yellow to golden hair (except in 7desa).
Mesonotum without conspicuous white pollinosity, appearing metallic from almost
any viewpoint . 2

— Hair of pleural regions Pees Hair of epeequacsal ridge black (cope in rb
Hair of fore coxae entirely black or at most with a few yellow hairs near base.
Mesonotum of ¢ with entirely black hair. Mesonotum with rather thick white
pollinosity on anterior part of prescutum, usually over humeral calli and usually
also on a supra-alar spot each side of scutum (pollinosity usually conspicuous to
naked eye, but if not then conspicuous under microscope from some viewpoint

as fly is turned) . 7
Genal hair blackish brown. Mesonotal hair of 3 blackish brown. c ponies
with surstyli and cerci as in Text-fig. 77 : R. idesa

[This species is known only from Walker’s ¢ Roane on an fae
Australian locality but appears to be distinct from any of the subsequently
described nominal species. ]

Genal hair yellowish white to golden yellow. Mesonotal hair of g mainly or entirely

pale yellow to golden or golden orange. surstyli and cerci not so shaped c 3
Species from Kei Islands. Mesopleuron with well developed white pollinose spot.
Abdominal Tr + 2 with all black hair . - : 5 - . ‘ R. nana

REVISIONARY CLASSIFICATION OF RUTILIINI 63

Species from Australia. Mesopleuron without white pollinose spot and brilliant
metallic, or at most with very thin pollinose overlay only visible at certain angles.
Abdominal Tr + 2 with some pale yellow hair at least on mid-venter and often
also on anteroventral part, sometimes on whole ventral surface : 4

Ground colour of parafrontals conspicuously darker than that of ne a
brown to blackish; inner margins of parafrontals (along the lines of frontal setae)
usually with distinct traces of metallic green showing through the pollinosity
(especially in 2), and each side of vertex extensively metallic green or blue-green
between eye and ocellar triangle. Wing base of § strikingly explanate, the wings
appearing to have prominent basal ‘shoulders’. Third antennal segment
mainly dark brown. Abdomen of ¢ appearing to naked eye semipellucid tawny
reddish with silvery blue or blue-green reflections (i.e. not with clear cut bands on
a dark ground). g surstyli and cerci as in Text-fig. 81. : : . R. formosa

Ground colour of parafrontals reddish yellow to reddish and therefore not noticeably
contrasting with ground colour of parafacials; inner edges of parafrontals without
trace of metallic colour, at most only slight trace of metallic greenish showing on
vertex. Wing base of § not explanate. Third antennal segment bright orange
(except usually some reddish brown suffusion in panthea). g abdomen usually not
so coloured, with medially interrupted transverse green bands on dark ground
colour. ¢ surstyli and cerci not exactly of this shape (though similar). . : 5
Vestiture of fore coxae and hypopleura entirely golden orange. Setulae of post-
ocular row, the vertical setae, and some thoracic setae usually golden red.
Abdominal T5 with some pale yellow hair laterally. Parafrontal hair usually
largely pale yellow or whitish (sometimes all dark, especially in 2). Third antennal
segment unicolorous orange. Abdomen with the dark hind band of T3 slightly
bowed forwards on each side, so that the transverse green band of the tergite is
narrower near the middle on each side than elsewhere. More elongate cee
in which the abdomen is conspicuously longer than its width . 2 6
Vestiture of fore coxae and hypopleura partly black (the strongest setae black).
Setulae of postocular row, vertical and all thoracic setae black. Hair of abdominal
T5 entirely black. Parafrontal hair all blackish. Third antennal segment
partly suffused with reddish brown colour, especially on outer side. Abdomen
with dark hind margin of T3 forming a transverse band of even width (no definite
bowing forwards on each side), the metallic band of this tergite therefore also of
even width across the abdomen. More robust species with abdomen not notice-
ably much longer than its width . - : R. panthea
Basicosta pale yellow. Setae of postalar callus ad fee at eal base golden red.
Median abdominal dark line evanescent, virtually absent on T4. [Western
Australia] . : : 2 . R. goerlingiana
[This nominal species is peceae not Semitic iia from caesia.]
Basicosta blackish brown (if slightly pale then at least dark brown on anterior edge).
Setae of postalar callus and fringe of costal base black (latter may have a very few
red setulae intermixed). A fine purplish black median line distinct along all
abdominal tergites. [Western Australia (including Monte Bello Is.) through
Northern Territory to North Queensland] E : ' . RR. caesia
Suprasquamal hair orange-yellow. Mesopleuron shining ieillianthy, without white
pollinose spot. Thorax without very thick white pollinosity over humeral calli
and each side of scutum. ¢ surstyli as in 2 i 78. {Oriental Region to
Queensland] , ‘ : R. rubriceps
Suprasquamal hair black. pie pledin — cise ‘siiite peliniose spot or
extensively coated with white pollinosity conspicuous in some light (except in
corona). Thorax with thick white pollinosity over humeral calli and usually on
each side of scutum (if not evident to naked eye then distinct in some lights

64

Io

II

I2

13

R. W. CROSSKEY

under microscope). ¢ surstyli and cerci usually differently sais gr very

similar in chersipho and caeruleata) : 8
Ground colour of parafrontals dark brown to Pagan ome fread pers e dee

to naked eye). Genal hair blackish brown. Third antennal segment dark brown

to brownish black (at most only reddish orange at extreme base). Abdomen of g

usually showing some tawny brown colouring to naked eye visible — the

metallic colouring . 9
Ground colour of parafrontals jallen to reddish oeiare (head es distantly

yellow to naked eye). Genal hair yellow. Third antennal segment bright

orange or yellow-orange (slightly suffused with darker reddish colouring in

occasional specimen or brownish in cvyptica). Abdomen of 3 usually not pgs

very evident tawny colouring through the metallic colour : 10
¢ genitalia with surstyli and cerci as in Text-fig. 75; surstylus with a isbn oars

nence submedially on the anterior edge and cerci in oe slender with the apical

part bent slightly forwards . : 5 : R. imperialis
6 genitalia with surstyli and cerci as in Text: fg. 76; _ surstyius rather slender and

without submedian prominence on anterior edge, cerci in profile rather bulbous

medially with the apical part more or less straight and narrowed abruptly from

the bulbous middle part : C R. imperialoides sp. n. (p. 67)
Mesonotum black or purple-black with a “bold ‘pattern of silver-blue markings as

follows: pair of lines submedially on prescutum reaching to suture, spot over

humeral callus, supra-alar spot on scutum, spot on hind part of notopleuron,

small spot on outer edge of prescutum just mesad of notopleuron (all spots with

white pollinosity and very shifting appearance). Scutellum not metallic, very dark

reddish brown (almost blackish brown to naked eye); posteromedian part of

scutum with dark red-brown colouring similar to scutellum when seen under micro-

scope (but black to naked eye). Abdomen blackish brown with very strongly con-

trasting spots or bands of light metallic silvery blue or silvery green; the bands or

spots widely separated medially and shape of those on T3 and T4 sublunate.

[Western Australia; g surstylus as in Text-fig. 84] . - s R. chersipho
Mesonotum without such pattern, ground colour not black but ranging from bright

brassy green to deep blue-violet, usually with distinct darker vittae (dark

specimens of caeruleata with deep blue-violet mesonotum may show pair of paler

blue prescutal lines but if so then scutellum metallic violet also). Scutellum

metallic, colour ranging from green to violet. Abdominal bands or spots deep

golden green to violet-blue (rarely pale silvery in caeruleata but then scutellum

violet); bands or spots of intermediate tergites usually not at alllunate . c II
Upper two-thirds or more of postorbits silvery white pollinose, sharply contrasting
in colour with the golden yellow genae and postbuccae. [Eastern Australia] . 12

All of the postorbits distinctly yellow pollinose, colour of postorbits therefore not
strongly contrasting with the golden yellow genae and agate re [Western
Australia] . : 15

Mesopleuron with distinet white pollinose spot which § is se conspicuous to ‘naked
eye. Body length usually 14-18 mm (occasional ¢ specimens ares smaller).

Cerci and surstyli of g of varied form . 13

Mesopleuron without white spot, entirely setae. Tengen I es mm. ¢ surstyli
and cerci in profile similar to vubriceps (Text-fig. ee surstyli in posterior view
similar to splendida (Text-fig. 72) . : ; . R. corona

3d genitalia with cerci and surstyli as in Text- fie, 74: See of extraordinary
form, with large forwardly directed tooth-like process and strongly acuminate
between this process and the apex (the large tooth projecting laterally when
hypopygium viewed from behind) ; apical part of cerci rather straight and slender
or at most slightly sinuous, the tip not bending forwards or backwards. Lower
calypter haired on most of its surface (occasional specimen with hairing only in

REVISIONARY CLASSIFICATION OF RUTILIINI 65

basal depression of calypter). Calyptrae not or only extremely faintly infuscate,

basal part of upper calypter not unusually opaque white. Mesonotum with very
broad bold continuous black sublateral vittae : : R. cryptica sp. n. (p. 65)

— 6 genitalia otherwise shaped, surstylus without such tooth-like process and cerci

either with tips bent forwards or backwards (Text-figs 72 & 73). Lower calypter

with surface hairing mainly confined to basal depression and area adjacent to

suprasquamal ridge. Calyptrae distinctly infuscate on at least the apical half,

the basal half of the upper calypter unusually opaque white (the white colour very

conspicuous to naked eye on side view of fly and strongly contrasting with the

smoky lower calypter). Sublateral dark vittae of mesonotum either bold,
broad and continuous or evanescent on scutum and broken at the suture . : 14

14 6 genitalia with surstyli and cerci as in Text-fig. 73; surstylus with blunt antero-

median projection and the cerci is profile very strongly recurved at their apices.

Sublateral black vittae of mesonotum very broad, bold and continuous (no
exceptions seen) . : ‘ . R. decora

— 6 genitalia with surstyli and cerci as in Texte fig. 72: suratylus simple, without

trace of prominence on its anterior edge, and cerci rather straight for most of their

length but with the extreme tips bent distinctly forwards. Sublateral black

vittae of mesonotum sometimes very broad, bold and continuous (especially in

specimens from southern part of range) but usually fine and inconspicuous,

evanescent on scutum and broken at transverse suture in Queensland specimens
R. splendida

15 Mesonotum dark bluish green to deep blue-violet, with the darker vittae very

inconspicuous to naked eye (pair of lighter blwe lines sometimes present on

prescutum). Surstylus of $ genitalia in profile as in Text-fig. 79, not deeply

concave on its anterior edge. White aati of thoracic dorsum very
inconspicuous to naked eye . ; R. caeruleata

— Mesonotum pale brassy green with the four black! vittae vexy boldly marked.

Surstylus of $ genitalia in profile rather deeply concave on anterior edge and

strongly acuminate on apical half (Text-fig. 82). White pollinosity of thoracic dor-

sum conspicuous to naked eye on prescutum anteriorly between the vittae, on
humeral callus and on supra-alar area of scutum - ; : R. transversa

Rutilia (Chrysorutilia) cryptica sp. n.
(Text-fig. 74)

6. Head. Ground colour mainly yellow-orange to reddish orange on parafrontals and
interfrontal area, genal dilations with some slight metallic golden green reflections in some
lights; pollinosity mainly bright yellow, thinner on parafrontals than on parafacials, but
pollinosity of postorbits silvery white over dark ground colour; upper occiput dark metallic
green; parafrontals with a little dark brown colouring just below vertex. Parafrontal and
most of parafacial hair black or brownish black; genal, postbuccal and lower parafacial hair
yellow to golden orange; occipital hair yellowish white. Vertex 0:21-0:24 of head-width (0-22
in holotype). Frontal setae very fine, mostly hair-like, lowermost ones crossing at tips. Gena
048-061 of eye-height (0-56 in holotype). Parafacial very wide, about 4-0 times as wide as
third antennal segment, completely haired. Antennae reddish orange on first two segments
and base of third, remainder of third segment suffused with darker reddish brown to brown
colouring; antennae falling short of epistome by about their own length; third segment about
2-1 times as long as second segment. Palpi yellow. Thorax. Dorsum golden green to viola-
ceous blue with four very broad bold black vittae and blackish side-margins, the submedian
pair of vittae stopping on scutum just beyond transverse suture, the sublateral black vittae
continuous to hind margin of scutum and neither broken nor constricted at transverse suture;
white pollinosity present over humeral calli, anteriorly on prescutum over the metallic areas
between the pairs of black vittae, and on supra-alar spots, the pollinosity only conspicuous in

66 R. W. CROSSKEY

some lights. Sides of thorax purplish black or very dark brownish black with metallic areas
coloured blue-green to purple-violet on upper sternopleuron and mesopleuron, the mesopleural
metallic area largely covered by a bold white pollinose spot (conspicuous to naked eye) ; metallic
areas of sternopleuron and mesopleuron often outlined with more violaceous colour than rest of
metallic area. Thoracic hair (including suprasquamal ridge hair) entirely black. Wéngs.
Basicosta black. Wing membrane clear hyaline (except for usual brown sub-basal area).
First basal cell almost completely devoid of microtrichia. Calyptrae semi-translucent whitish,
lower calypter at most only very faintly smoky, upper calypter not strikingly opaque white on
basal part and therefore not very conspicuous to naked eye in side view; lower calypter with
long dark erect hair on most of its surface (occasional specimen with hair confined to basal
depression of lower calypter). Legs. Black with all black hair. Hind tibia with the ad fringe
well developed and without pd setae. Abdomen. Ground colour blackish, purplish black or
very dark tawny brownish with a pattern of golden green, light bluish green or slightly violaceous
metallic areas arranged in spots or bands; black median vitta distinct to naked eye. T1 + 2
with a long transverse metallic area, slightly diffuse on each side; T3 with a pair of submedian
and a pair of lateral metallic spots, the submedian and lateral spot of each side sometimes nar-
rowly or broadly coalesced so that each side of T3 has a metallic band; T4 with a pair of sub-
median and a pair of lateral metallic spots which appear always to be well separated; T5 with a
large metallic area on each side which is slightly excavate on its anterior margin so that the
metallic area is slightly V-shaped or slightly lunate. All abdominal hair black, recumbent
except on last tergite. Genitalia (Text-fig. 74): surstylus of very remarkable form, with a large
sub-basal to submedian anteriorly and outwardly directed process and strongly excavate
between this process and the apex, apical part as a whole strongly acuminate (the anterior
process conspicuous in posterior view of hypopygium as well as in profile); cerci very slender
on the apical third and this part very slightly sinuous, but actual apices of cerci not directed
either forwards or backwards. Measurements. Body length about 11-5-15 mm, wing length
about 12°5-15°5 mm.

Q. Generally similar to ¢ but ground colour of abdomen more distinctly black and metallic
areas of abdomen tending to be more coppery green. White pollinosity of mesonotum more
conspicuous than in g and very conspicuous to naked eye between the black vittae of the pre-
scutum and laterad of the sublateral vittae. No proclinate orbital setae. Vertex 0:27-0:29
of head-width. Hind tibia with either one or two small pd setae. Probably averaging larger
than ¢ (three speciments seen with body length about 15 mm).

MATERIAL EXAMINED

Holotype g, AUSTRALIA: South Australia, near Moonta, 16.ix.1904 (W. Wesche).
In British Museum (Natural History), London.

Paratypes. AUSTRALIA: 2 g, Victoria, Bright (HW. W. Davey). 1 g, Victoria
(C. French). 13, New South Wales, Barrington Top, 13-17.xii.1g21 (G. Goldfinch).
23,39, ‘Australia’ (no other data) (ex coll. Bigot). All paratypes in British Museum
(Natural History).

DISTRIBUTION. Known only from south-eastern Australia.

AFFINITIES. R. (C.) cryptica sp. n. was in the past confused with R. (C.) splendida
(Donovan) and R. (C.) decora to which it is certainly extremely closely allied. It has
very similar or in some specimens identical metallic colour patterning to these species,
but is distinguished from both of them (and from all other Rutiliini) by the quite
exceptional shape of the surstyli (Text-fig. 74) mentioned in the description and the
key; the surstylus shape is so characteristic that the species can be recognized
immediately the male genitalia are examined. The shape of the apical part of the
genital cerci also distinguishes cryptica from decora and splendida (in cryptica the

REVISIONARY CLASSIFICATION OF RUTILIINI 67

tips of the cerci are neither bent forwards nor backwards, whereas in decora they
are recurved and in sflendida bent forwards at the extreme tips). Other features
_ are sufficiently well correlated with the genital differences to be of some value in
distinguishing cryptica from splendida and decora; these include the non-infuscate
lower calypter (lower calypter extensively smoky brown in sflendida and decora)
and the unusually hairy lower calypter (in nearly all specimens of cryptica the lower
calpter is haired on most of its surface, whereas in the other two species the lower
_ calypter is hairy only inits basal depression). R. cryptica is of interest in this charac-
_ ter because the extent and density of long dark erect hairing is much greater than
in any other species of the subgenus Chrysorutilia (and therefore the lower calyptrae
_ are much more hairy than in any other Rutiliine, as lower calyptral hairing is found
_ only in the segregate Chrysorutilia and nowhere else in the tribe Rutiliini).

Rutilia (Chrysorutilia) imperialoides sp. n.
(Text-fig. 76)

3g. Head. Ground colour of parafacials and genae mainly brownish orange, epistome more
tawny yellow, parafrontals and outer margins of parafacials against the eyes blackish brown;
genal dilations with golden green reflections in some lights (especially towards postbuccae) ;
upper occiput dark metallic green; pollinosity mainly whitish over hind parts of genae and along
_ parts of parafacials against the eyes, brownish yellow on upper anterior parts of genae and on
parafacials (except against eye margins), greyish yellow on parafrontals; postorbits greyish
white pollinose over dark ground; interfrontal area brick red. Hair of parafrontals, para-
facials, genae and postbuccae black; occipital hair yellowish white. Vertex 0-15-0-17 of head-
' width. Frontal setae irregular, hair-like, apices of rows not meeting. Gena 0:36-0:40 of
_ eye-height. Parafacial very wide, about 4:5 times as wide as third antennal segment, entirely
haired. Antennae very small, falling short of epistomal margin by much more than their
' own length; basal segments dark reddish brown (more reddish orange on apex of segment 2),
_ third segment brownish black except for narrow trace of orange colour at junction with second
_ segment; third segment about twice as long as second segment. Palpireddish orange. Thoraw.
| Mesonotum dark metallic green with variable tinges of blue and violet, and with four broad
bold black vittae; the submedian pair of vittae stopping on scutum just behind transverse
suture, the sublateral pair appearing to naked eye to be completely continuous but actually
_ narrowly broken at transverse suture (the prescutal part of the sublateral vitta stopping just
_ short of transverse suture so that prescutum is very narrowly metallic green between end of
the black line and the suture itself); humeral calli and anterior metallic parts of prescutum
with distinct overlay of white pollinosity, supra-alar area only with an exceedingly thin trace
of whitish pollinosity visible in some lights (i.e. no definite white pollinose supra-alar spots).
_ Scutellum metallic dark green to violet). Sides of thorax black with metallic dark green to
' violaceous area on upper sternopleuron and on disc of mesopleuron; the mesopleural metallic

area overlaid by white spot of thick pollinosity (most conspicuous from above). Thoracic

hair (including that of suprasquamal ridge) entirely black. Wings. Basicosta black. Wing

membrane clear hyaline (except for usual brown sub-basal mark). First basal cell with micro-

trichia along its length. Calyptrae yellowish white with some very faint trace of yellowish brown

suffusion. Lower calypter almost completely bare, only a few erect dark hairs at extreme

base of basal depression adjacent to suprasquamal ridge. Legs. Black with black hair. Hind

tibia with ad fringe short, especially on apical third, and without pd setae or with one minute

submedian fd setula. Abdomen. Dark reddish brown with a broad black conspicuous median
_ vitta and with metallic areas on all tergites which range from cupreous to blue-green. Metallic

areas arranged as follows: transverse strip on each side of Tr + 2; band on each side of T3

68 R. W. CROSSKEY

which occupies the anterior two-thirds of tergite length; similar but narrower band on each side
of T4 occupying only about half tergite length; large metallic area on each side of the black
median vitta on T5 which occupies most of side of tergite and extends forwards to abut directly
against hind margin of preceding T4. All abdominal hair black; dorsal hair recumbent on
Tr + 2 to T4 (at most a little erect hair anteromedially on T3 and hairing of hind margin of
T4 semi-erect as usual). Genitalia (Text-fig. 76): Surstylus in profile rather slender and simple
(without trace of process on anterior edge), cerci in profile conspicuously bulbous medially
and rather abruptly narrowing into the long, slender and rather straight apical part. Measure-
ments. Large species, body length about 16-17 mm, wing length about 15-16 mm.

(2. Not positively associated. Specimens in BMNH collection believed to be probably
females of imperialoides have following features: Generally darker than g with dark greenish
colour of mesonotum less conspicuous because of stronger development of thick white pollinosity
combined with very heavy black vittae; white pollinosity thick and conspicuous over humeral
calli, supra-alar spots, three longitudinal lines on prescutum (between and on outer side of
the submedian black vittae), over notopleuron and on a small spot on extreme outer edge of
prescutum. Calyptrae more infuscate than in g, opaque white base of upper calypter there-
fore more conspicuous. Abdomen very black with deep gold-green or cupreous metallic spots
or areas as follows: transverse strip on each side of T1 + 2; pair of submedian and pair of
lateral spots on T3; pair of very small (much smaller than those on T3) submedian spots and
pair of lateral spots on T4; very large subtriangular metallic area on each side of T5 which
does not reach forwards as far as the hind margin of T4 (i.e. separated from latter by a narrow
strip of black ground colour). Metallic spots on each side of T3 sometimes partially or com-
pletely coalesced into a transverse band across each side of tergite. No proclinate orbital setae.
Vertex 0-26 of head-width. Hind tibia with one or two small pd setae. Averaging larger than
6, body length about 17-19 mm, wing length about 16-18 mm.]

MATERIAL EXAMINED

Holotype g, AuSTRALIA: New South Wales, Wee Jasper, xii.1g20. In British
Museum (Natural History), London.

Paratypes. AUSTRALIA: Ig, same dataas holotype. 1, Victoria (F. du Boulay).
Both paratypes in British Museum (Natural History).

In addition to the g holotype and paratypes cited above I have seen four female
specimens in the BMNH collection which probably are conspecific. However, as
positive association of these females with the males cannot be made at this time I
exclude them from the type-series. Their data are: 1 9, Victoria, Monbulk, 1895;
I 9, Victoria (C. French) ; 1 9, New South Wales, Katoomba, Blue Mts, 3000-3300 ft,
1.1912 (Dodd junior); and 1 9, Australian Capital Territory, Paddy’s River, 2.ii.1955
(Paramonov).

DISTRIBUTION. South-eastern Australia including Victoria, New South Wales
and Australian Capital Territory.

AFFINITIES. During the present revisionary work it was found that two quite
distinct forms of genitalia were found amongst males of the supposedly single species
previously identified by authors as Rutilia imperialis Guérin-Méneville. In one
form (Text-fig. 75) the surstylus has a blunt submedian prominence on the anterior
edge and the cerci in profile are very slender and have the apical part bent slightly
forwards; in the other form (Text-fig. 76) the surstylus is much narrower and
completely lacks any trace of a prominence on the anterior edge, and the cerci in
profile are very distinctly bulbous medially with a rather sudden contraction to the
straight slender apical part. This difference can be recognized immediately the

REVISIONARY CLASSIFICATION OF RUTILIINI 69

male genitalia are examined (after removal from the fly), and there appear to be no
intermediates. It has therefore been concluded that ‘imperialis’ of authors consists
of a confusion of two very closely allied species which are indistinguishable on
external characters (or apparently so as none have been found of any value). As
Guérin-Méneville’s type-material (which included a male specimen) is lost it was
uncertain to which of the two species his name imperialis should be applied; this has
therefore been resolved by neotype designation (see p. 125). The species with the
prominence on the surstylus and the very slender cerci appears to be more common
than the other species and the neotype specimen chosen for imperialis (which shows
the genital characters clearly although the hypopygium remains in situ) is therefore
of this species; there is no available name for the second species, which is therefore
here described as new with the name imperialoides. The new species is clearly
extremely closely allied to imperialis and examination of male genitalia is essential
for accurate identification of imperialoides and imperialis.

Subgenus DONOVANIUS Enderlein stat. n.

Donovanius Enderlein, 1936 : 409. Type-species: Musca regalis Guérin-Méneville, 1831, by
original designation.

Psavonia Enderlein, 1936 : 414. Type-species: Psaronia bisetosa Enderlein, 1936, by original
designation. Syn. n.

Menevillea Enderlein, 1936 : 416. Type-species: Rutilia pellucens Macquart, 1846, by original
designation [presumed correctly identified, but see discussion]. Syn. n.

Diacnosis. Parafrontals pollinose, not metallic. Epistome and genae non-metallic. Facial
carina flattened, slightly widened at level of base of third antennal segment, merging into lunula
without a distinct depression. Parafacials bare or partially haired (hairing if present normally
not reaching as low as bottom of eye). 9 normally without proclinate orbital setae (one or
two present in some specimens). Arista micropubescent. Humeral callus with 3-4 setae,
inner one or two sometimes scarcely differentiated from hair, very raiely only outer two present.
Posthumeral setae absent or one inner posthumeral developed just mesad of humeral callus.
Usually one post ia seta, occasionally none, rarely a small second seta developed. Scutum
with or without some supernumerary prescutellar setae. Postalar callus with 4 (rarely 5)
strong setae. Suprasquamal ridge thickly long haired. Scutellum flattened; with 4-10 pairs
of marginal setae (these sometimes, stiff, straight, slightly spiniform); marginals not preceded
by any preapical setae. Hair of lower part of pteropleuron not developed in front of level of
posterior sternopleural seta. One sternopleural seta (o + 1), rarely trace of a small anterior
stpl seta (especially in 9). Prosternum bare; prosternal membrane with or without some long
soft hair. Hind tibia with long regular anterodorsal fringe (without evident ad setae amongst
it) and without fd setae or with only one such seta (very rarely trace of second fd). Abdomen
with median depression in last tergite (T5). T3 without median marginal setae (rarely one pair
in 9), with lateral marginal setae. T5 with median transverse row of strong erect setae.
Sternite 5 of § with simple non-prominent rounded lobes. ¢ genitalia with distal membranous
part of distiphallus subequal in length to or at most only slightly longer than sclerotized proximal
part, and with extremely large foliaceous surstyli (Text-figs 66-71). [Mainly large forms mea-
suring 15-23 mm, with dark head ground colour and usually dark unicolorous body, never
with bold spot pattern.]

DISTRIBUTION. Mainly distributed throughout Australia and Tasmania, but a
few species also in Solomon Islands, New Hebrides, Fiji and Samoa. Unknown
from the Oriental Region and New Guinea.

70 R. W. CROSSKEY

Discussion. This subgenus includes nearly one-third of the species of Rutilia s.1.,
and contains most of the large brown, blackish or purplish or dark green Australian
species in which, even to the naked eye, the end of the abdomen appears excavate
because of the median depression or groove in the last tergite. As early as 1775
Fabricius had described a species of this subgenus with a name (R. retusa) which
apparently alludes to the blunt-ended appearance given to these flies by the apical
excavation, and several common eastern Australian species belonging in Donovanius
were described by the other early authors, such as imusta Wiedemann, regalis
Guérin-Méneville, vividinigra Macquart and sabrata Walker.

One of the species undoubtedly belonging in this subgenus is R. pellucens Macquart,
for which a neotype is designated in this paper. This species was cited by Enderlein
(1936) as the type-species of his genus Menevillea, but there are some slight discrepan-
cies between the characters cited by Enderlein for Menevillea and those shown by
the true pellucens as identified by Macquart and fixed by neotype (for example
Enderlein mentions the presence of marginal setae on T3 and ‘Discalmacrochaeten’,
i.e. preapicals, on the scutellum). I have not seen the two female specimens deter-
mined, and cited as pellucens by Enderlein (1936 : 416), but for the present am pre-
suming Enderlein’s identification to be correct, in which case it follows that Mene-
villea is a new synonym of Donovanius Enderlein. However, if it should prove
(when Enderlein’s specimens are located) that his pellucens was misidentified, then
the name Menevillea would fall as a synonym of Grapholostylum or just possibly
Rutilia s.str., but in any event it is certainly a synonym of an older name. (It is
therefore of no practical importance whether Enderlein identified pellucens correctly
or not.)

Enderlein’s genus Psavonia was characterized mainly by having a single pair of
median marginal setae on T3 and by little else that notably distinguished it from
Donovanius. It was based only on two female specimens (one herein designated
lectotype). Examination of the lectotype of the type-species, bisetosa, shows that
there are really no differences which justify holding Psaronia as a distinct taxon
from Donovanius at supraspecific level and Psaronia is therefore placed as a synonym
of the latter name.

Reference needs to be made here to Enderlein’s genus Psaroniella, for which Rutiha
castanipes Bigot was cited as type-species. In this case it is known positively that
Enderlein misidentified the type-species, for I have seen the single female specimen
(from Victoria, Koonwarra, Gippsland) that Enderlein cited as castanipes and
found that it belongs to a completely different species from that described by Bigot.
The true castanipes Bigot (type-material in BMNH examined) is a species of the
subgenus Donovanius here defined, and the name is a junior synonym of R. inusta
(Wiedemann) ; the specimen misidentified by Enderlein as castanipes actually belongs
in the subgenus Rutilia s.str., and is a specimen of R. setosa Macquart. The generic
name Psaroniella Enderlein is therefore a synonym of Rutilia s.str. and not of
Donovantus.

The affinities of subgenus Donovanius seem clearly to lie most closely with Chryso-
rutilia. Both subgenera have four or more postalar setae, both have a very well
developed fringe or comb of close-set setulae on the anterodorsal surface of the hind

REVISIONARY CLASSIFICATION OF RUTILIINI 71

tibia, both have extremely bushy and often crinkled hair on the suprasquamal
ridge and there is normally only a single (posterior) sternopleural seta; other
resemblances include the occurrence in most forms of hairing on the prosternal
membrane, and general similarity in the shape of the facial carina (which tends to be
rather flattened on the anterior face and to merge rather gradually into the lunula
so that there is only a weak depression where carina and lunula coalesce). The
broad sulcate last abdominal tergite with its transverse row of strong setae provides
the most obvious external feature distinguishing Donovanius from Chrysorutilia,
though the difference in the pteropleural hairing mentioned in the subgeneric key
and shown in Text-figs 19 & 20 appears to apply constantly throughout the sub-
genera and to provide a real distinction. The large heavy foliaceous surstyli of the J
genitalia of Donovanius are very different from the narrow pointed surstyli found in
Chrysorutilia (cf. Text-figs 66-71 & 72-84), and the g genitalia therefore readily
distinguish these two subgenera. Another minor difference between them, but one
which is apparently constant, is the lack of any hairing actually on the prosternum in
Donovanius which contrasts with the presence of at least a few fine hairs on the
anterior corners of the prosternum in Chrysorutilia.

The species of Rutilia s.str. look superficially much like Donovanius on account
_ of the grooved apex to the abdomen, but are at once separable by the presence of
only three postalar setae and by the lack of a hind tibial fringe, and by the differently
shaped ¢ surstyli (cf. Text-figs 54-57 & 66-71).

Donovanius species appear to be entirely unrepresented in the Oriental Region,
but the distribution of the subgenus is more extensive in the Pacific islands than that
of other subgenera, which are either confined to Australia (Neorutilia, Ameniamima,
Rutilia s.str.) or else do not occur so far is known anywhere further east than New
Guinea and Australia (Grapholostylum) or the New Hebrides (Chrysorutilia). One
species of Donovanius is known from Fiji (tvansfuga Bezzi) and two from Samoa
(savatiensis Malloch and nigrihivta Malloch), whence these are, respectively, the only
Rutiliines known to occur. Samoa in Polynesia represents the easternmost limit
_ of distribution of the tribe Rutiliini as a whole.

The British Museum (Natural History) collection contains a female specimen of
the Samoan species Rutilia (Donovanius) nigrihirta Malloch which was reared from a
larva of a Lucanid beetle identified as a species of Aegus Macleay, probably A.
wpoluensis Arrow; this seems to be the first fully authenticated host record for a
species of subgenus Donovanius.

The species of Donovanius do not aggregate in any obvious way, and no species-
_ groups are recognized within the subgenus. The ¢ genitalia with their very distinc-
| tive form of surstyli are extraordinarily uniform in all the species (or supposed species)
of the subgenus.

Most of the synonymies shown in the list of included species that follows require
no special comment, but amplification is needed here concerning R. (D.) sabrata
(Walker) and R. (D.) bisetosa (Enderlein). One of the species of Donovanius is a
large blackish brown form with dark violet reflections and unusually long antennae
compared to those of other Rutilia species, and it was to this species that Malloch
(1927 : 347; 1929 : 300) applied Guérin-Méneville’s name inornata, using (in the

72 R. W. CROSSKEY

later paper cited) the antennal length as a key character; as a result, Paramonov
following Malloch has identified some specimens of the long-antenna species in
different museum collections as tmornata Guérin-Méneville. Neither Malloch nor
Paramonov saw Guérin-Méneville’s two original syntypes in Paris Museum, and
Malloch’s identification—working from the description alone—was reasonable;
however, examination of the two inornata types during the present work (one of
them bearing an original Guérin-Méneville label and herein designated lectotype,
p. 72) has shown that the true znornata belongs in the subgenus Rutilia s.str. and
is the same species as Guérin-Méneville’s desvotdyi (=vivipara Fabricius). There
are some discrepancies between the original description of inornata and the charac-
ters shown by the lectotype, especially as the legs are not as black as the description
implies, but I see no reason to doubt that the two specimens in Paris are the two that
Guérin-Méneville mentioned. Hence it is concluded that Malloch misidentified
inornata. But the species that he called by this name showing the long antennae
was described by Walker with the name sabrata, and this is the valid name for the
species.

From comparison of Enderlein’s types of bisetosa (described from the 9 in Psaronia)
and migribasis (described from gf and 9 asa variety of Donovanius fulgidus (Macquart))
I am convinced that the respective 9 and ¢ lectotypes designated in this paper are
conspecific and I therefore place nigribasis as a synonym of bisetosa. The name
bisetosa is chosen to stand valid for the species in preference to migribasis because it
alludes to the character of a pair of median marginal setae on the third (apparent
second) abdominal tergite which are present in both sexes, and because bisetosa is the
name upon which Enderlein’s so-called genus Psaronia was based. It should be
added that the 9 lectotype of bisetosa and the g lectotype of nigribasis have the same
type-locality, namely Marloo Station, Wurarga, Western Australia, which strongly
supports the conclusion that both names apply to the same species.

INCLUDED SPECIES

Rutilia (Donovanius) agalmiodes (Enderlein) comb. n. AUSTRALIA (Queens-
land). [Holotype examined].
R. (D.) analoga Macquart. AUSTRALIA (Victoria to Queensland). [Holotype
examined].
dubitata Malloch syn. n. [Holotype examined].
R. (D.) bisetosa (Enderlein) comb. n. AUSTRALIA (Western Australia). [Lecto-
type examined].
mgribasis (Enderlein) syn. n. [Lectotype examined].
R. (D.) brunneipennis Crosskey sp. n. SoLomon IsLANDs (Guadalcanal).
[Holotype examined].
R. (D.) ethoda (Walker). AusTRALIA (Western Australia). [Holotype exa-
mined].
R. (D.) inusta (Wiedemann). AUSTRALIA (all states). [Lectotype examined].
castanifrons Bigot syn. n. [Holotype examined].
castanipes Bigot synn. [Lectotype examined].

REVISIONARY CLASSIFICATION OF RUTILIINI 73

potina (Walker) syn. n. [Holotype examined].
spinipectus Thomson syn. n. [Lectotype examined].
R. (D.) lepida Guérin-Méneville. Austratia (A.C.T., New South Wales,
Victoria). [Neotype examined].
fulgida Macquart syn. n. [Lectotype examined].
onoba (Walker) syn. n. [Holotype examined].
R. (D.) nigrihirta Malloch. Samoa (Upolu). [Holotype examined].
R. (D.) pellucens Macquart. AvusTRALia (A.C.T., New South Wales, Victoria).
[Neotype examined].
imitator (Enderlein) syn. n. [Holotype examined].
R. (D.) regalis Guérin-Méneville. Austratta (A.C.T., New South Wales, South
Australia, Victoria). [Neotype examined].
nigra Macquart (nomen nudum).
R. (D.) retusa (Fabricius). AUSTRALIA (Western Australia). [Holotype
examined].
aditha (Walker) syn. n. [Holotype examined].
viriditestacea Macquart syn. n. [Lectotype examined].
R. (D.) sabrata (Walker). AusTRALIA (New South Wales, Queensland). [Holo-
type examined].
[inornata Guérin-Méneville sensu authors. Misidentification. ]
R. (D.) savaiiensis Malloch. Samoa (Savaii). [Holotype examined].
R. (D.) spinolae Rondani. AusTRALiA. [Type-material not located, possibly
lost; species unrecognized, tentatively assigned to Donovanius].
R. (D.) transfuga Bezzi. Fiji (Viwa, Viti Levu). New Hesripes (Eromanga,
Espiritu Santo, Malekula, Tana). [Holotype examined].
R. (D.) viridinigra Macquart. AusTRALIA (New South Wales, Queensland).
[Lectotype examined].
barcha (Walker) syn. n. [Holotype examined].
fuscotestacea Macquart syn. n. [Holotype examined].

Kery TO SPECIES OF THE SUBGENUS DONOVANIUS

[Note : The limits of species in this subgenus are very difficult to determine. There are few
structural characters that appear to have much value in separating species, and the § genitalia
are so uniform (or at least differ in such a subtle and intangible way) that they have little value
for species recognition. It is often difficult reliably to associate males and females, and there
is clearly much variation within a species in such features as hairiness of parafacials, develop-
_ment of median marginal setae on T3, and the number of proclinate orbital setae in females.
At present the entities considered to be species differ mainly in their general appearance as
shown by body colour, hair colour and size; these features seem to separate specimens into
fairly convincing species, but there is considerable intergradation (new synonymies established
_ above are based on absolute agreement between types). The key here given must be treated
| as very tentative: it is by no means certain that the named taxa represent distinct species,
and museum collections may contain undetermined specimens that will not conform with the
key. R. spinolae is omitted as the type has not been located and the name remains enigmatic. ]

1 Australian species . : : : : : ; : : ; : : 2
— Melanesian and Samoan species : 5 : ; ¢ : : : - 12

74

R. W. CROSSKEY

Pleural regions of thorax with pale yellow or golden hair (in bisetosa mesopleural
hair mainly dark and some dark hairs on SRE Hair of suprasquamal
ridge yellow : 3

Pleural regions of thorax wich black an (occasionally some inconspicuous pale yelloe:
hair on fore margin of mesopleuron, on barette and posteroventral part of ptero-
pleuron). Hair of suprasquamal ridge black or brownish black (some ae
of sabvata with some yellow hair intermixed with dark hair) . 6

Wing base explanate, wings appearing to have strongly developed al ‘shoulders’.
Colouring distinctive, prescutum and scutum dark green, scutellum violet, abdo-
men light tawny yellow with sharp black median line. Abdomen with entire
venter of Tr +2 and mid-venter of T3 pale yellow haired. Surstyli of 5
genitalia longer and narrower than usual (Text-fig. 65)

R. agalmiodes (3 only, 2? unknown)

Wing base not noticeably explanate. Not so coloured, if abdomen rather light
tawny then mesonotum only very dull greenish and scutellum not violet.
Abdomen almost entirely black haired (except in analoga). ¢ surstyli of usual
large wide form (e.g. as in Text-fig. 67) . : : ; : - : 4

Hair of abdominal T5 pale yellow or golden. Parafrontal hair of g¢ mainly
yellowish white (black in 9). Mesonotum dull dark greenish or greenish brown
or sometimes blackish brown in 9, with conspicuous whitish pollinosity, slightly
contrasting with the tawny (g) or red-brown (?) abdomen; abdomen with distinct
dark centre line. Parafacials bare or partially haired . . R. analoga

Hair of abdominal T5 black. Parafrontal hair black in both sexes. Galea varied
but not as above, whitish pollinosity of mesonotum very inconspicuous. Para-
facials more or less completely haired . 5

Pleural hair entirely yellow. First basal oe of ee ete microtrichial 2
Abdominal T3 without median marginal setae. eae generally dark,
slightly metallic, greenish blue or violaceous . : : . RR. retusa

Pleural hair mainly blackish on mesopleuron and Pomretienes on upper edge of
sternopleuron. First basal cell with microtrichia.2 Abdominal T3 usually with
a pair of median marginal setae. Colour varying from dark bronze-green to
coppery brown or blackish brown, sometimes with purplish red tinge on abdomen

R. bisetosa

Predominantly green species, colour ranging from light golden green to dark green
or blue-green. Abdomen with fine dark median line and dark tergite hind
margins well visible to naked eye . . yy

Colour not green but ranging from red-brown to piaeke Bega or Ree blac
sometimes with bronze or coppery purple tinges (rarely slight trace of very dark
green colour visible but then over an almost black general colour). Dark median
abdominal line and dark tergite hind margins inconspicuous to naked eye (except
in § of pellucens) : 8

Abdomen of g with ground poloue ae eee mete: (T3 ane T4) pale ‘cme
reddish, the tawny colour clearly visible to the naked eye through the metallic green
or bluish tints (especially as fly is turned); these two tergites also with rather
thick white overlay of pollinosity which makes them appear distinctly white when
viewed from behind. ¢ usually 15-17 mm in body length 5 . . R. lepida

(Q not definitely associated: golden green to blue-green females resembling
those of vegalis but with elongate third antennal segment (c. 4—5 times as long as
second segment) appear to be Jepida]

Abdomen of $ without such appearance, ground colour all dark and intermediate
tergites with only very thin inconspicuous whitish pollinosity basally, abdomen
therefore appearing a pure golden green to blue-green (occasional specimen
coppery green). ¢ usually 17-19 mm in body ‘enyth : : : . R. regalis

{Females of golden green to blue-green colour with antennae of normal length

12

13

REVISIONARY CLASSIFICATION OF RUTILIINI 75

(third antennal segment about 3 times as long as second segment) are almost
certainly regalis]
First basal cell without microtrichia.2_ Blackish species with a purple or violaceous
tinge in most specimens, occasionally a dark greenish black tinge (some dark
brownish black specimens without metallic tinge occur and resemble inusta:
careful check on presence or absence of microtrichia along first basal cell essential
on such specimens) : : 9
First basal cell with gxiecoicialaia alenn its male length (at least eden. Red-
brown to very dark blackish brown species, seaeailel without sitesi tinge nee
coppery purple tinge in ethoda) . 10
Antennae unusually long, falling short of (eetistemaicll macemiss by shea as noch as
their own length in g and by conspicuously less than their own length in 9; third
segment about 3:5 times as long as second segment in g and 4-5 times as long
as second segment in ? (facial carina correspondingly elongate, especially in 9).
Thorax with some very pale yellow hair on anterior edge of mesopleuron, on
ventral edge of humeral callus, around prostigmatic area, on barette and postero-
ventrally on pteropleuron (sometimes also some pale hairs on suprasquamal
ridge). Apical pair of scutellar setae conspicuously smaller than other scutellar
marginal setae . : R. sabrata
Antennae of normal small size, esnally falling short of pistol margin by more than
their own length in both sexes; third segment about 2-2-5 times as long as second
segment in both sexes (facial carina not unusually elongate). Thorax without
any pale hair. Apical scutellar setae ae not conspicuously smaller than
other scutellar marginal setae - : . RR. viridinigra
Thorax and abdomen with burnished coppery bronze or purplish reflections, some-
times also traces of green colouring. [Western Australia] : ; . R. ethoda
Body without such burnished reflections (at most only the scutellum faintly violet).
{Eastern Australia from Tasmania to Queensland] . : II
Abdomen of $ light reddish brown or tawny brown with conspicuous black median
line, abdomen of 2 blackish brown or almost black with the median line scarcely
detectable to naked eye. Smaller species, length usually 13-16 mm R. pellucens
Abdomen of both sexes very dark, brownish black to black, sometimes very dark
reddish brown on T3 in the g§ (but ¢ abdomen consistently darker than the more
tawny abdomen of $ pellucens). Larger species, length usually 16-20 mm_—_—*iR«. inusta
{Note that the females of znusta and pellucens are not reliably distinguishable
on present evidence]
Body unicolorous dark purplish brown. Epistome blackish. All hair black.
Wings heavily infuscate, especially on anterior half. Calyptrae blackish brown.
Very large species, length about 20 mm. [Solomon Islands]
R. brunneipennis sp. n. (p. 76)
Body either unicolorous green or with green to blackish mesonotum and mainly
tawny or reddish brown abdomen. Ground colour of epistome reddish yellow.
Hair black or yellow hair present on pleural regions. Wings not infuscate or (in
nigrihivtia) with yellowish brown staining along veins. Small to medium sized

species, length 10-16 mm. [New Hebrides, Fiji or Samoa] : 13
Hair entirely black. Legs black. Colour uniformly dark green on “thorax and
abdomen. [Samoa] . : ‘ R. nigrihirta

Hair of pleural regions of thorax and most of ‘the fore comal hair pale yellow to
golden orange. Legs partly reddish yellow. Abdomen mainly tawny yellow to

2 The microtrichia of the first basal cell are very small, and care is needed to determine whether the
cell is bare (i.e. without microtrichia beyond the brown-pigmented base) or whether it has a band of
microtrichia along its length (even when present the microtrichia are sometimes confined to an area
along the middle of the cell). The cell needs to be examined by transmitted light at a magnification
not less than x 100.

76 R. Wi CROSSE.

reddish brown and contrasting with green or blackish mesonotum (some specimens

with golden green abdomen rather similar to mesonotum but then pale ground

showing through the green colour). . 14
14 Unusually small species, length 1o-11-5 mm. Bristling of eater of Steraoplemen

mid and hind coxae and prescutellar region not noticeably spiniform. Femora

entirely reddish yellow. [Samoa] . : R. savaiiensis
— Larger species, length 12-5-16 mm. Bristling of verter of Srecouplcnes mid and

hind coxae, prescutellar region and scutellum very strong, stiff, rather straight

and sub-spiniform. Femora either reddish yellow or partly or mostly darkened

to reddish brown or dark brown. [Fijiand New Hebrides] . - R. transfuga

Rutilia (Donovanius) brunneipennis sp. n.

9. Head. Ground colour brownish black on most of head, genal regions dark reddish brown,
epistome black. Head pollinosity very thin and whitish, bases of parafrontal hairs appearing
as slightly darkened minute dots. Parafrontal and parafacial hair black, genal hair blackish
brown, occipital hair yellowish white. Vertex 0:25 of head-width. Frontal setae very fine,
meeting or crossing at tips. Proclinate orbital setae absent. Gena very wide, 0-50 of eye-
height. Parafacial about three times as wide as third antennal segment, with short sparse
black hairs on most of its height. Antennae black and slightly elongate (facial carina corres-
pondingly slightly longer than usual), third segment about 2:5 times as long as the rather long
second segment; antennae falling short of epistomal margin by about 1-3 times their length.
Palpi brownish with tawny apices. Thorax. Dark purplish brown with no evident metallic
colour to naked eye (mesonotum with slight coppery purple glints under microscope) ; mesonotum
without obvious pollinosity and without definite dark vittae (thin trace of whitish pollinosity
present on prescutum and visible under microscope when seen from behind). All thoracic
hair black. Scutellum with eight pairs of marginal setae (apicals included), the apical pair
very much smaller than other marginals; scutellar marginals (and also the prescutellar setae)
rather stiff and straight. o-+ 1 stpl setae. Wings. Basicosta black. Wing membrane
distinctly infuscate, especially anterobasally, suffused with dark brown along the veins and
especially on v—m cross-vein. First basal cell completely microtrichiate, as the rest of the
wing. Calyptrae blackish brown with brown fringe hair. Legs. Black with entirely black
hair. Hind tibia with well formed even close-set ad fringe. Abdomen. Dark purplish brown
with entirely black hair, concolorous with thorax. T3 without median marginal setae. Hair
of T3 recumbent, of T4 semi-erect, and of T5 erect. Measurements. Large species, body
length about 20 mm, wing length about 19 mm.

6. Unknown.

MATERIAL EXAMINED
Holotype 2, SoLomon IsLanps: Guadalcanal, Suta, 27.vi.1956 (E. S. Brown).
In British Museum (Natural History), London.

DISTRIBUTION. Known only from the holotype from Guadalcanal in southern
Solomon Islands.

AFFINITIES. R. brunneipennis sp.n. unquestionably belongs in the subgenus
Donovanius, although it should be noted that the holotype is aberrant in one respect:
the postalar callus of the left side has the normal 4 strong setae, whereas that of the
right side has only three postalar setae. It is the only species of the subgenus yet
known from the Solomon Islands. The heavy infuscation of the wings (to which the
specific name alludes) makes the species appear quite distinctive, though in the
general very dark and uniform colouring it is rather similar to R. (D.) inusta, to
which it is perhaps closely allied. The g is unknown, but as the ¢ genitalia in

REVISIONARY CLASSIFICATION OF RUTILIINI 77

Donovanius are of very little or no use for identification it is considered justified to
describe this new species from a female holotype.

Subgenus RUTILIA Robineau-Desvoidy

Rutilia Robineau-Desvoidy, 1830 : 319. Type-species: Tachina vivipara Fabricius, 1805, by
subsequent designation of Crosskey (1967 : 26).

| Psaroniella Enderlein, 1936: 417. Type-species: Rutilia castanipes Bigot sensu Enderlein
[misidentification] [= Rutilia setosa Macquart, 1847], by original designation. Syn. n.

Stivaulax Enderlein, 1936 : 428. Type-species: Tachina vivipara Fabricius, 1805, by original
designation. [Isogenotypic name with Rutilia Robineau-Desvoidy, junior objective synonymy
first noted by Crosskey (1967 : 26)].

_ Dracnosis. Parafrontals pollinose, not metallic. Epistome and genae non-metallic. Facial

carina with anterior surface of upper part convex or with slight median ridge and only lower
part flattened on anterior surface, sides slightly pinched-in ventrally, separated from lunula
| by distinct depression well visible in profile. Parafacials bare or haired. 2 with proclinate
| orbital setae (normally one pair well developed, rarely two each side, very rarely absent).
| Arista micropubescent. Humeral callus with 3-4 setae. Posthumeral setae distinct in both
sexes, variably from 1-3 each side. One or two post ia setae. Scutum without supernumerary
prescutellar setae (rarely the last post acy seta doubled). Postalar callus with three strong setae.
Suprasquamal ridge haired or bare. Scutellum with slight dorsal flattening or hollowing just
before tip; with 5-7 pairs of marginal setae (these rather stiff and straight) ; with a well developed
transverse row of small preapical setae preceding the marginals. Pteropleuron not haired in
_ front of level of posterior sternopleural seta. Two or three sternopleural setae (1 + I or 2 + 1).
_ Prosternum and prosternal membrane bare. Hind tibia without definite anterodorsal fringe
or with short inconspicuous fringe, normally from 1—3 ad setae well developed, 2—4 well developed
_ pd setae or occasionally one pd only ing. Last abdominal tergite (T5) with median depression
(rather small and shallow in confusa). T3 with transverse row of several (usually 6-12, but
_ often only 2-4 in confusa) strong erect often spiniform median marginal setae, the row slightly
| arcuate forwards so that middle setae of the transverse row are more distant from hind margin
_ of the tergite than the others; T3 also with lateral marginal setae. T5 with median transverse
| row of strong erect setae. Surstyli of J genitalia of rather varied form (Text-figs 54-57). [Dull
| reddish brown, tawny or blackish brown forms, with little or no metallic colouring, at most
| with very slight reddish violet or greenish tinge dorsally on thorax and intermediate abdominal
tergites].

DIsTRIBUTION. Occurring only in Australia from Tasmania to Queensland.

Discussion. Before considering the characteristics and interrelationships of
_ Rutilia in the strict sense it is necessary briefly to discuss the type-species of the
_ genus and its fixation, for there has been some confusion in past taxonomic bibliog-
raphy; although, fortunately, this has not affected the generic concept of Rutilia.
_ The genus when originally described by Robineau-Desvoidy contained four nominal
' species, one of which was Rutilia vivipara (Fabricius), which Fabricius (1805) had
' described in the genus Tachina Meigen. Robineau-Desvoidy’s (1830) identification of
| vivipara related to a specimen that stood in the collection of Count Dejean, and may
or may not have been correct, but there has never been any means of confirming the
rightness of Robineau-Desvoidy’s identification because both the specimen from

|

| Dejean’s collection and Fabricius’ type of vivipara are lost. But there are a few
discrepancies between the brief descriptions of Fabricius and Robineau-Desvoidy
(the latter for instance mentioning a bluish tinge on the mesonotum) and on the basis

78 R. W. CROSSKEY

of these Guérin-Méneville (1843 : 264) concluded that Robineau-Desvoidy was
dealing with a different species from Fabricius (though as he had not, it seems certain,
seen the specimens of either author his conclusion was based on little more than
guesswork). Following upon his conclusion, Guérin-Méneville (op. cit. : 264, 269)
described the species supposedly misidentified by Robineau-Desvoidy as Rutilia
desvoidy Guérin-Méneville, and subsequent references to the Rutiliini contain various
citations of the type-species of Rutilia as vivipara or as desvoidy1. I have earlier
(Crosskey, 1967 : 26) shown that none of these citations were valid for type-fixation,
either because the fixation was made ambiguously, or the name cited was not that
of a nominal species or was that of a species not originally included in Rutilia, and I
therefore published a designation of Tachina vivipara Fabricius as type-species of
Rutilia. Inthe present revisionary work it is now most desirable to establish beyond
any future doubt the identity of vivipara Fabricius, as this species is the nomen-
clatorial pivot of the whole Rutiliini, and to show that desvoidy: Guérin-Méneville
is in reality a junior synonym of vivipara Fabricius.

The original description of vivipara might not be readily accessible to Australian
dipterists likely to work on the Rutiliini, and I therefore quote it here: Tachina
vivipara

‘T. pilosa grisea, abdomine ferrugineo, scutello cupreo.

Habitat in Insulis maris pacifici carnes consumens vivipara. Dom. Billardiere.

Magna. Caput griseum, ore albido. Thorax pilosus, griseus, fusco lineatus.
Scutellum cupreum, nitidum. Abdomen pilis atris, rigidis hirtum,
subferrugineum, segmentorum marginibus nigris. Pedes pallide testacei.’

All later workers have assumed, and I agree with this assumption, that the
provenance of Fabricius’ specimen was almost certainly Australia, though recorded
as ‘Insulis maris pacifici’; localities of origin were often very imprecisely recorded at
the time when Fabricius was writing, and although some Rutilia are now known
from the Pacific islands it cannot be said that Fabricius’ description fits any of them
at all well. On the other hand, for such an early work, it is an extremely good
description of a common Rutilia species which occurs throughout eastern Australia
from Cape York to Tasmania and which has been commonly identified as vivipara
for many years. This species has entirely reddish yellow legs, a ferruginous colour
often with rather coppery scutellum, conspicuous blackish brown thoracic vittae,
blackish hind margins to the abdominal segments, and exceptionally strong rather
spiniform abdominal bristling, and is therefore a virtually perfect match with
Fabricius’ statements (respectively “‘pedes pallide testacei’, ‘abdomine ferrugineo’,
‘scutellum cupreum, nitidum’, ‘thorax . . . fusco lineatus’, “abdomen . . . segmen-
torum marginibus nigris’, and ‘rigidis hirtum’). There is only one large (Fabricius’
‘magna’) species of Rutilia with all pale legs, strongly vittate thorax, dark reddish
brown colouring, and such strongly bristled abdomen, and it is to this species that
Fabricius’ name vivipara unquestionably applies and it is from this species that a
neotype for vivipara has been designated elsewhere in this paper (see p. 126).

With the identity of vivipara Fabricius objectively pinned down by neotype it is
now possible to determine whether desvoidyi Guérin-Méneville is distinct from vivi-
para or not. One of three original syntypes of desvoidyi still exists in the Paris

REVISIONARY CLASSIFICATION OF RUTILIINI 79

Museum and is herein designated as lectotype (see p. 119); the lectotype is in rather
poor condition (some mould, holes in body, a mid and a fore leg missing) but it
shows the bare parafacials, three postalar setae, haired suprasquamal ridge, entirely
reddish yellow legs, brown abdomen with darkened hind margins to the segments,
yellowish pleural hair, and same form of chaetotaxy, and is considered undoubtedly
conspecific with the neotype of vivipara. Hence desvoidyi Guérin-Méneville is here
placed as a new synonym of vivipara Fabricius, and it is concluded that vivipara of
Robineau-Desvoidy (1830) (=desvoidyi according to Guérin-Méneville) is actually
therefore the same species as vivipara Fabricius and not a misidentification as
Guérin-Méneville supposed. Consequently there is no longer any confusion over the
type-species of Rutilia: Tachina vivipara Fabricius is an originally included nominal
species, correctly identified by Robineau-Desvoidy, and fixed as type-species of the
genus by subsequent designation of Crosskey (1967 : 26).

Tachina vivipara Fabricius is also type-species by original designation of Stivaulax
Enderlein, and this name thus falls as a junior objective synonym of Rutilia Robineau-
Desvoidy. One other name enters into junior synonymy with Ruizlia s.str., namely
Psaromella Enderlein, the type-species of which was cited by Enderlein (1936 : 417)
as castanipes Bigot; but Enderlein misidentified Rutilia castanipes Bigot (which
belongs in subgenus Donovanius, the name a synonym of imusta Wiedemann) and
the single specimen from Gippsland, Victoria, that he cited belongs to Rutilia (Rutilia)
setosa Macquart, and the generic name Psaroniella is therefore a synonym of Rutilia
s.str. and not of Donovanius. (The 9 specimen identified as castanipes by Enderlein
is in MNHU collection, Berlin, and has been examined: it is labelled ‘Koonwarra
Gippsland, Victoria’ and has a determination label in Enderlein’s hand reading
‘Psaroniella castanipes (Big. 1880) 9 Dr Enderlein det. 1936’.)

Rutilia s.str. contains only a small number of species, and although the typical
group of the genus in a nomenclatorial sense is not very representative of the wide
range of forms included in the genus as a whole. The rather strongly developed
ventral marginal bristling of the abdominal tergites, especially in vivipara itself,
sets the subgenus rather apart from all the other Ritilia, and the possession of only
three strong setae on the postalar callus makes the included forms rather obviously
different from superficially similar large brown forms with depressed tip to the
abdomen found in the subgenus Donovanius. In vivipara the abdominal chaetotaxy
is more strongly spiniform than in all other Rutilia s.1. and the marginal setae of the
ventral ends of the tergites are so strong and stiff that they simulate the similar very
strong setae found in Formosia; but whereas in Formoszia the bristles of the tergite
venters are directed almost straight downwards those of R. vivipara are directed
backwards or mainly so (and those of the female are shorter and more stubby than
those of the male). The third abdominal tergite in Rutilia s.str. has unusually well
developed median marginal setae, which often enable specimens of the subgenus to
be distinguished at once from other subgenera. Normally the T3 median marginals
form a transverse row of about six to a dozen erect setae, the row bowing forwards
near the centre so that the middle one or two pairs of marginal bristles are not close
to the hind edge of the tergite like the rest (and are therefore less truly marginal) ;

80 R. W. CROSSKEY

in other subgenera median marginal setae are normally absent on T3 or represented
by only a single fine pair.

Rutilia s.str. is most easily distinguished from Chrysorutilia, Donovanius and
Ameniamima, by the possession of only three instead of four or more setae on the
postalar callus, and from Chrysorutilia and Ameniamima by the absence of ptero-
pleural hairing in front of the posterior sternopleural seta. It differs also from
Chrysorutilia and Donovanius by lacking a long hind tibial fringe (though a short one
is present in confusa), and usually by having two distinct pd setae on the hind tibia.
The depression in the last abdominal tergite, and the presence of two or three strong
sternopleural setae, also, separate Rutilia s.str. from Chrysorutilia.

In species of Rutilza s.str. the suprasquamal ridge may be bare or haired (though
presence or absence of such hairing seems to be constant within each species). In
vivipara the ridge is haired, but the hairing is rather short and sparse (as in the
species of Grapholostylum) instead of very long, dense and crinkly (as in the species
of Donovanius, Chrysorutilia, and Neorutilia), a fact which perhaps suggests that the
affinities of Rutilia s.str. lie more closely with Grapholostylum than with any other
subgenus, a supposition which is supported by the several other character that
Rutilia and Grapholostylum share in common (among them the three postalars, lack
of hind tibial fringe, unusually strong development of the chaetotaxy of the thoracic
dorsum, pteropleuron bare in front of the posterior st¢p/ seta, and rather similar form
of facial carina).

If all the species of Rutilia s.str. and Grapholostylum are considered it is found that
there are one or two species which are rather intermediate and tend to form an
interconnecting link between the two subgenera, as discussed in more detail under
Grapholostylum. One of these, here placed in Rutilia s.str., is the aptly named
species confusa Malloch. This species has the suprasquamal ridge bare and on this
account was originally described by Malloch as a Formosia, but it has none of the
characters of true Formosia apart from the bare ridge and is certainly a Rutilia in
all other respects (just as the species of Ameniamina subgen.n. with bare supra-
squamal ridge are equally Rutilia in the wide sense) ; but confuwsa has only some 2-4
(rarely more) median marginal setae on abdominal T3, has a somewhat shallow
median depression in T5, and has a definite though short hind tibial fringe, and so
differs in these features from vivipara; on the other hand, it has rather strong ventral
marginal setae on the tergites, colouring very like that of vivipava, and the form of
5th abdominal sternite of the male, which all confirm the correct placement of
confusa in Rutilia s.str.

The following points should be noted about the synonymies indicated in the list
of included species. The neotype specimen herein designated for Rutilia durvillet
Robineau-Desvoidy (see p. 124) is conspecific with the neotype of vivipara, and
durvillei therefore goes into synonymy; apart from being justified by what little is
known of durvillei from the original description it is desirable to dispose of this name
as a synonym since it has never been in use for a recognized species. Enderlein
(1936 : 430) suggested that durvillet was perhaps only a variety of desvoidyi (=vivi-
para) ; the present fixation of the neotype eliminates the name completely as a junior
synonym. For Rutilia inornata Guérin-Méneville an original syntype specimen

REVISIONARY CLASSIFICATION OF RUTILIINI 81

still exists in Paris Museum and is here designated as lectotype (see p. IIg); it has
three postalar setae, haired suprasquamal ridge, and all the other features shown by
vivipara and is believed to be undoubtedly conspecific with the neotype of vivipara;
hence inornata also is placed as a synonym of vivipara. (But note that the name
inornata has been misapplied in the literature to the species that should be called
sabrata Walker and belongs in Donovanius: see further discussion of this under that
subgenus. )

INCLUDED SPECIES

Rutilia (Rutilia) confusa (Malloch). AusTRaALia (A.C.T., New South Wales,
South Australia, Victoria). [Holotype examined].
R. (R.) dentata Crosskey sp. n. AUSTRALIA (Victoria). [Holotype examined].
R. (R.) setosa Macquart. AUSTRALIA (New South Wales, Victoria). [Neotype
examined].
[castanipes Bigot sensu Enderlein, 1936. Misidentification.}
R. (R.) vivipara (Fabricius). AUSTRALIA (A.C.T., New South Wales, Queens-
land, Tasmania, Victoria). [Neotype examined].
desvoidyi Guérin-Méneville syn. n. [Lectotype examined].
durvillei Robineau-Desvoidy syn. n. [Neotype examined].
inornata Guérin-Méneville syn. n. [Lectotype examined].

KEY TO THE SPECIES OF SUBGENUS RUTILIA

1 Suprasquamal ridge haired : . ; : : : : : R. vivipara
— Suprasquamal ridge bare . 2
2 Legs entirely reddish yellow. Seactasty, especially in 3, swith a dark green tinge to

naked eye and with rather inconspicuous presutural vittae. Surstylus of ¢
genitalia shaped as in Text-fig. 55 . : R. confusa
— Legs partially dark brown to blackish brown. Scutum not appearing dark green to
naked eye, sometimes with coppery violet tinge. Surstylus of $ genitalia not of
this shape . 3
3 Parafacials entirely haired. Tarsi reddish ‘yellow to reddish and concolorous with
tibiae. Surstylus of g genitalia broad basally with an anteromedian toothlike
process, deeply excavate between this process and apex (Text-fig. 56)
R. dentata sp. n. (p. 81)
— Parafacials bare or at most only haired at extreme upperends. Tarsi blackish, much
darker than tibiae. Surstylus of f awrer ae and ered tapering to sharp
apex (Text-fig. 57) : : : . &. setosa

Rutilia (Rutilia) dentata sp. n.
(Text-fig. 56)

[Specific name alludes to a tooth-like process on ¢ surstylus. |

6. Head. Ground colour dark brownish or blackish on occiput and genal dilations and
/parafrontals, light tawny brownish or reddish on other parts; interfrontal area red-brown;
pollinosity yellowish white. Parafrontal hair black, parafacial hair light golden orange (with a
few dark hairs intermixed at extreme upper end of parafacial), genal hair pale yellow, occipital
hair yellowish white. Vertex 0-13 of head-width. Frontal setae not, or only just, meeting at
apices. Gena 0-35-0:39 of eye-height. Parafacial wide, about 3-8 times as wide as third

82 R. W. CROSSKEY

antennal segment, completely haired. Antennae brownish orange basally, third segment
dark brown except for some bright orange-red colour at junction with second segment, falling
short of mouth-margin by twice the length of the third segment; third segment about 2-1 times
as long as second segment. Palpitawny brown. Thorax. Ground colour brownish or black-
ish, a little reddish on sides around the sutures of the pleurites; dorsum with dark coppery
pink tinge to naked eye, and with traces of greenish yellow reflections under microscope
(especially along transverse suture); trace of greenish reflection on sternopleura. All dorsal
hair black; hair of pleural regions light yellow to pale golden orange, except for some blackish
hairs on the upper and posterior parts of the mesopleura; hair also blackish on sides of humeral
calli. Scutellum with 6-7 pairs of marginal setae (apicals included), the apicals as strong as the
other marginals. Thoracic chaetotaxy not noticeably spiniform. Wangs. SBasicosta black.
Wing membrane nearly clear hyaline, entirely microtrichiate. Calyptrae semi-translucent
pale brownish, with pale yellow hair fringe. Legs. Reddish or reddish yellow except for some
brown to blackish brown darkening on coxae and basally on femora, tarsi concolorous with
tibiae. Most of the coxal hair and the long soft posterior hair of the fore and mid femora, also
the long hair of the basal anterior surface of the hind femora, pale yellowish; other leg hair black,
except for the usual reddish hair on the a and av surfaces of the fore tibia. Chaetotaxy of mid
and hind tibiae variable: mid tibia either without or with one or two minute ad setae; hind tibia
with short regular inconspicuous ad fringe (with or without a distinct ad seta submedially in the
fringe), and with one or two fd setae. Claws long. Abdomen. Slightly paler than thorax,
dark reddish brown with some darkening of hind margins of intermediate tergites (which appear
slightly blackish to naked eye), and with a narrow blackish median line; intermediate tergites
with slight coppery purple reflections and with traces of coppery green reflections in some lights;
these tergites largely overlaid with very thin whitish pollinosity which is hardly at all visible
to naked eye. All abdominal hair black except for a few pale yellowish hairs at extreme base
of venter; long hairs and setae of T5 with reddish apices. T3 with a row of many median
marginal setae, these rather short and slightly stubby. Dorsal hair of Tr + 2 and T3 recum-
bent, that of T4 semi-recumbent basally but erect distally, hair of T5 very long and fine and
entirely erect. Sternite 5 with each lobe very slightly excavate on apical margin. Genitalia
with cerci and surstyli as in Text-fig. 56; surstylus of unusual form, very broad basally and with
a blunt prong or tooth-like process on anterior edge submedially, excavate between this tooth
and the apex, and bearing some extraordinary multifid setae on inner surface directed inwards.
Measurements. Large species, body length about 16-18 mm, wing length about 15-17 mm.

©. Generally similar to gj except that the abdomen is much more black and has the whitish
pollinosity of the intermediate tergites much more conspicuous, purplish reflections less notice-
able. Some pale yellow hair present on notopleural swellings and on extreme sides of scutum
(below supra-alar setae). rons with one pair of well developed proclinate orbital setae.
Vertex 0:23-0:25 of head-width. Size asingd.

MATERIAL EXAMINED
Holotype g, AUSTRALIA: Victoria, Monbulk. In British Museum (Natural

History), London.

Paratypes. AUSTRALIA: I g, 2 9, same data as holotype (one 9 with year date
‘1895’ on label in addition to words ‘Monbulk Victoria’). 1 2, Victoria, Grampians,
Reed’s Lookout, 23.xii.1953 (B. McMullan). All paratypes in British Museum
(Natural History).

DISTRIBUTION. Known only from south-eastern Australia.

AFFINITIES. Closely allied to other species of Rutilia s.str. and perhaps most
closely to R. (R.) setosa Macquart, from which it differs most notably in the key
characters cited above. The genitalia, with their remarkable form of surstyli, are

REVISIONARY CLASSIFICATION OF RUTILIINI 83

most distinctive, and no other Rutiliine is known in which a surstylus of this shape
occurs (Text-fig. 56) or in which the genitalia have strong setae with divided apices.

Subgenus GRAPHOLOSTYLUM Macquart stat. n.

Grapholostylum Macquart, 1851 : 196 (223). Type-species: Grapholostylum dorsomaculatum
Macquart, 1851, by monotypy.

Agalmia Enderlein, 1936 : 433. Type-species: Rutilia albopicta Thomson, 1869 [=Grapholo-
stylum dorsomaculatum Macquart, 1851], by original designation. Syn.n. [Junior homonym
preoccupied by Agalmia Enderlein, 1934 (Muscidae)].

Diacnosis. Parafrontals pollinose, not metallic. Epistome and genae not metallic. Facial
carina slightly flattened to strongly convex on anterior surface, outline convex in profile and
sharply distinguished from lunula; epistome prominent. Parafacials bare or haired. @ pro-
clinate orbital setae usually absent, at most one pair (sometimes present one side and absent
the other). Arista long-pubescent to short-plumose. Humeral callus with 4 setae, usually
well developed. Posthumeral setae one or two. One very strong post ia seta [one specimen
seen with small second fost ia]. Scutum without supernumerary prescutellar setae. Postalar
callus with three strong setae. Suprasquamal ridge haired. Scutellum convex dorsally without
flattening towards apex; 4-6 pairs of marginal setae; with row of well developed preapical
setae before the marginals. Pteropleuron not haired in front of level of posterior sternopleural
seta. Two strong sternopleural setae (1 + 1) [one specimen seen with small second anterior
stpl]. Prosternum bare; prosternal membrane bare or sparsely haired (variable in same species).
Hind tibia without anterodorsal fringe, usually two distinct ad setae and two or three pd setae.
Last abdominal tergite without median depression, evenly convex across its width and sides
strongly tapering posteriorly (hypopygium prominent and well visible in profile). T3 without
median marginal setae or with one pair or with a transverse row of semi-erect small median
marginals developed to variable extent; T3 with lateral marginal setae. T5 with some long
fine erect discal setae (often irregular in § but tending to form transverse preapical row in 9).
Sternite 5 of ¢ acuminate posterolaterally and with a submedian pair of conspicuous downwardly
directed prominences (Text-fig. 33), these visible in situ in profile. ¢ genitalia with apical
membranous part of distiphallus whip-like, exceptionally long and slender (about twice as long
as sclerotized basal part of distiphallus: Text-fig. 38), surstyli long and narrow and cerci longer
than surstyli (Text-fig. 64). [Medium-sized to large species with white pollinose spots over
mesopleura and sternopleura and often with white pollinose spots distinguishable over humeral
calli to notopleura and in supra-alar areas].

DistRIBuTION. Eastern Australia from Tasmania to Queensland.

Discussion. Gyvapholostylum was originally described by Macquart (1851) as a
genus allied to Rutilia having one included species, viz. G. dorsomaculatum. For
many years the identity of Macquart’s genus remained enigmatic, but Townsend
(1932 : 38; 1936 : 153; 1938 : 416) rightly showed that Grapholostylum is a true
Rutiliine. Regrettably, Enderlein (1936 : 441), guessing from the description of
Macquart and overlooking Townsend’s (1932) paper, decided that the genus belonged
to the Ameniinae and placed it near Amenia Robineau-Desvoidy—a rather bad case
of misidentification, which was corrected by Crosskey (1965 : 103-106), who showed
that the misidentified G. dorsomaculatum sensu Enderlein is actually Amenia sex-
punctata Malloch. For the present work Macquart’s type-material of G. dorso-
maculatum (including the lectotype designated elsewhere: Crosskey (1971 : 271))
has been studied in detail and Townsend’s placement in the Rutiliini found to be
correct ; it should be emphasized, however, that the name applies to a valid species

84 R. W. CROSSKEY

and is not a synonym either of Rutilia decora Guérin-Méneville or of R. splendida
(Donovan) as Townsend stated (1932 : 38-39; 1938 : 416). On the other hand,
the name is a senior synonym of Rutilia albopicta Thomson (syn. n.), the type-species
designated by Enderlein (1936) for his genus Agalmia, and because of this specific
synonymy the generic name Agalmia Enderlein, 1936, is a new synonym of Grapholo-
stylum Macquart. (Here it should be noted that Agalmia of Enderlein (1936) in
the Rutiliini is a junior homonym of Agalmia Enderlein (1934) in the Muscidae; no
replacement name is needid as Agalmia Enderlein, 1936, is itself a synonym.)

Grapholostylum, as here redefined as a subgenus within Rutilia s.l., contains a
small number of species that are very distinctive in the ¢ fifth sternite and ¢
hypopygial characters but which do not differ very strikingly on their other charac-
ters from certain other Rutilia. Certainly on the features of head form, chaetotaxy,
aristal hairing, and abdominal shape the subgenus shares many characters in
common with the subgenus Microrutilia Townsend, and in the presence of a well
formed transverse row of median marginal setae on T3 there is a marked resemblance
between one of the species of Grapholostylum, viz. subtustomentosa, and the subgenus
Rutilia s.str. However, subtustomentosa has the § sternite 5 and hypopygial charac-
ters exactly as in dorsomaculata (I can find no real differences at all between the
genitalia of the two species), and subtustomentosa is assignable to subgenus
Grapholostylum and not to Ruizlia s.str.

Sternite 5 in the male in this subgenus is shaped as in Text-fig. 33; the sides of the
sternite are drawn out to sharp points and there is a pair of large blunt downwardly
directed submedian lobes with a deep narrow cleft between them. The submedian
prominences are easily visible on the abdomen when seen in situ and project con-
spicuously in profile; the hind margin of the sternite is clearly excavate between the
submedian prominence and the lateral extremity of the sternite. A ¢ sternite 5 of
this form occurs nowhere else in the Rutiliini, and therefore makes Grapholostylum
an especially distinctive segregate.

The aedeagus in Grapholostylum differs from that of all other subgeneric segregates
of Rutilia sl. The membranous distal section of the distiphallus is exceptionally
long and slender, rather whip-like (Text-fig. 38), and about twice as long as the
sclerotized proximal section (which is of normal length) ; in other Rutilia s.1. the distal
section of the distiphallus is either shorter than, or about subequal in length to, the
proximal section. Species of subgenus Grapholostylum have, in fact, the most
elongate form of aedeagus found in the Rutiliini. Another slight difference in the
aedeagus between Grapholostylum and other Rutilia subgenera lies in the shape of the
epiphallus: in Grapholostylum the epiphallus, seen in profile, is widest well beyond
the junction of the distiphallus (Text-fig. 38), whereas in other subgenera the
epiphallus is widest at its base (i.e. near the junction with the distiphallus) and
contracts thence towards its apex (Text-fig. 37).

Apart from the genital differences, Grapholostylum differs from Rutilia s.str. by
lacking a median dorsal excavation in the last visible atdominal tergite (T5) and by
having a convex scutellum which lacks any definite flattening before the apex.
Differences between Grapholostylum and Microrutilia are discussed under the latter
subgenus.

REVISIONARY CLASSIFICATION OF RUTILIINI 85

The type-species, R. (G.) dorsomaculata, of this subgenus has white pollinose
markings on thorax and abdomen (to which Macquart’s name refers), but in general
the spots are less bold and obvious than those found in the subgenus Ameniamima
(q.v.), though they are mainly developed at the same sites—in humeral, supra-alar,
mesopleural and sternopleural positions. The white spots give dorsomaculata and
albovirida (the latter is possibly not specifically distinct from the former) some
superficial resemblance to Ameniamima species, but Grapholostylum differs much
from Ameniamima on the genital features already noted and by having three
postalar setae (four in Ameniamima) and a haired suprasquamal ridge (bare in
Ameniamima).

The size and colouring in dorsomaculata are rather variable, females ranging
from bright green to dark blackish green with coppery tinges, and males having a
light tawny brown abdomen with broad black median vitta. All specimens, how-
ever, have the white pollinose markings on the thorax and less obvious white
pollinose areas (somewhat shifting in appearance) on the abdomen; the thoracic
pattern appears always to include a pair of small submedian white spots medially
on the scutum just in front of the scutellum (spots in this position do not occur in
Ameniamima but are found in some of the Ameniine Calliphorids of the genus
| Amenia Robineau-Desvoidy, which slightly resemble the brighter green specimens of
| Grapholostylum species). As a result of the variability, and because earlier authors
misunderstood Macquart’s species, dorsomaculata has several synonyms (which are
_ here newly established after examination of types). Malloch’s species Rutilia
albovirida, described from the female, is probably the same as dorsomaculata but at
present there is insufficient evidence to establish definite synonymy.

Rutilia micans Malloch belongs in this subgenus and is rather intermediate between
dorsomaculata and subtustomentosa in hair colour and degree of development of the
_ white pollinose spots (the pattern of these is identical to dorsomaculata but they are
' less conspicuous in micans).

_ INCLUDED SPECIES

Rutilia (Grapholostylum) albovirida Malloch. AvusTRALIA (Queensland).
[Holotype examined].
R. (G.) dorsomaculata (Macquart). AusTRALIA (New South Wales). ([Lecto-
type examined].
albopicta Thomson syn. n. [Holotype examined].
fuscisquama Malloch syn. n. [Holotype examined].
leucosticta Schiner syn. n. [Holotype examined].
variegata Bigot syn. n. [Lectotype examined].
R. (G.) micans Malloch. AvusTRa.iA (Australian Capital Territory, New South
Wales). [Holotype examined].
R. (G.) subtustomentosa Macquart. AUSTRALIA (Tasmania). [Holotype
examined].
velutina Bigot syn. n. [Lectotype examined].

KEY TO SPECIES OF THE SUBGENUS GRAPHOLOST YLUM
1 Entire vestiture of the fore coxae orange or golden red. Bristling of fore femora red.

86 R. W. CROSSKEY

Prostigmatic and propleural setae red. All mesopleural hair and pteropleural
vestiture yellow or golden orange. Scutum without submedian white spots.
Mesonotum extensively tinged with light mauve-pink colour. Abdomen of both
sexes with transverse row of many small median ane setae on T3. 9 abdomen
without white spots. [Tasmania] . ‘ . R. subtustomentosa
— Vestiture of fore coxae all black or with at least the main 1 bristling black. Bristling of
fore femora black. Prostigmatic and propleural setae black. Mesopleural hair
partly or completely black, pteropleural vestiture partly black (at least the
strongest hairs black). Scutum with a pair of submedian white pollinose spots (in
addition to supra-alar pair), sometimes faintly marked. Mesonotum not exten-
sively mauve-pink, sometimes coppery pink in prescutellar area. Abdomen
normally without or with only one pair of median marginal setae on T3 (occasional
specimen with row of four or five such setae developed). 9 abdomen with some
diffuse white pollinose spots, usually including a pair of submedian spots on each
intermediate tergite (sometimes only visible as fly is turned) ; 2
2 Hair of sternopleuron pale yellow. Hair of abdominal venter entizely pale yellow
(6) or pale yellow ventrally on Tr + 2 and T3 (9). Hair of coxae partly or mainly
orange or golden red. At least some of snakes hair yellow. [A.C.T. and
New South Wales] : . R. micans
— Hair of sternopleuron eaelyie or oie bea Hom of eatire abdoauaal venter
black (both sexes). Hair of coxae entirely or almost entirely black. All meso-
pleural hair black . : 3
3 Size smaller, length 10-12 mm. Mesouotam aan metallic eneadd: ceceea or poled
green, without noticeable darker vittae when seen by naked eye. Lower half of
postorbit bright golden orange or yellow and upper part silvery white, transition
of the colouring rather abrupt. [Queensland] . : : R. albovirida
[Only known from 9, possibly not specifically distinct fom dopsormannae
— Size larger, length 12-18 mm. Mesonotum not brilliantly shining, sometimes green
but more often very dark olive-green to greenish brown and with some darkish vittae
evident to naked eye. Upper half of postorbit pale yellowish and not strongly
contrasting in colour with yellow or orange-yellow lower half, if upper part rather
white then normally not abruptly contrasted in colour with lower part (transition
of colouring more gradual). [New South Wales] . : : R. dorsomaculata

Subgenus MICRORUTILIA Townsend

Microrutiia Townsend, 1915 : 23. Type-species: Rutilia minor Macquart, 1846, by original
designation.

Prosenostoma Townsend, 1932 : 39. Type-species: Senostoma flavipes Brauer & Bergenstamm
sensu Townsend (misidentification) [=Rutilia (Senostoma) hirticeps Malloch], by original
designation. Syn. n.

Eucompsa Enderlein, 1936: 400. Type-species: Rutilia minoy Macquart, 1846, by original
designation. [Isogenotypic synonym of Microrutilia Townsend and junior homonym pre-
occupied by Eucompsa Enderlein, 1922 (Tabanidae)].

Pogonagalmia Enderlein, 1936 : 435. Type-species: Rutilia (Senostoma) hirticepbs Malloch,
1929, by original designation. Syn. n.

[Senostoma sensu authors, not Macquart. Misidentification.]

Diacnosis. Parafrontals pollinose or almost completely so, at most only metallic at extreme
upper ends in 9. Epistome and genae not metallic. Facial carina convex on anterior surface,
and strongly convex and well marked off from lunula in profile, upper part often very bulbous;
epistome strongly nasute. Parafacials bare or haired. @ proclinate orbital setae in one or two
pairs, rarely absent. Arista long-pubescent to short-plumose. Humeral callus with 4 setae,
rather strong. One or two posthumeral setae. Normally two strong post ia setae, occasionally

REVISIONARY CLASSIFICATION OF RUTILIINI 87

only one ing’, sometimes small third seta in front of main pair. Scutum without supernumerary
prescutellar setae. Postalar callus with three strong setae. Suprasquamal ridge haired.
Scutellum convex, not at all flattened before apex; 4-5 pairs of marginal setae (occasional
specimen with three only on one side) ; with row of preapical setae in front of marginals. Ptero-
pleuron not haired in front of level of posterior sternopleural seta. Two or three sternopleural
setae (I + 10r2-+ 1). Prosternum bare; prosternal membrane normally bare, rarely one or
two hairs. Hind tibia with short fringe or with irregular sparse long fringe, with one main ad
seta if fringe close-set but otherwise with several distinct ad setae, with three pd setae (occasional-
ly more). Last abdominal tergite without median depression, evenly convex across its width
and sides strongly tapering posteriorly, hypopygium very prominent. T3 without median
marginals or with a few weakly developed; T3 with lateral marginal setae. T5 with long erect
discal setae. Sternite 5 of g downwardly prominent and conspicuous in profile, each lobe
angulate posterolaterally (Text-fig. 32) and very slightly concave on its hind margin. ¢ genitalia
with distal membranous part of distiphallus shorter than sclerotized proximal part, surstyli
broad basally and slightly tapering to blunt end (Text-figs 60-63), sharply pointed at tip in
nigviceps. [Small or very small species of coppery green, emerald, or blue-green colour in
which scutellum often violaceous and male often with tawny yellow abdomen showing black
median vitta and cupreous or green tinges to ground-colour].

DIsTRIBUTION. Eastern Australia from Tasmania to Queensland, probably also
Western Australia.

Discussion. This subgenus is aptly named for, as redefined here, it still contains
the smallest species of Rwutilia s.l., and R. (M.) minor—the type-species—is the
smallest of all Rutiliini (as little in some specimens as 6-7 mm long). Enderlein
(1936) erected the genus Eucompsa for R. minor, but evidently realized before final
publication of his paper that Townsend (1915) had already proposed the genus
Microrutilia for the same species: thus Eucompsa is proposed on p. 400 of Enderlein’s
(1936) work and promptly sunk into synonymy with Microrutilia on p. 415 of the
same work. Apart from being a junior isogenotypic synonym the name Eucompsa
Enderlein, 1936, is also a junior homonym of one of Enderlein’s own generic names,
being preoccupied by Enderlein’s (1922) use of Eucompsa in the Tabanidae. Even
for Enderlein it was unusually careless nomenclature to publish a junior objective
synonym and a junior homonym in the same name.

The type-species of Pogonagalmia Enderlein, namely R. hirticeps Malloch, differs

_ from typical Microrutilia only in having the parafacials haired and in slight shape

differences (considered specific only) in the g cerci and surstyli, and none of these

distinctions would justify separating hirticeps from Microrutilia; hirticeps is here

considered to be consubgeneric with minor and the genus-group name Pogonagalmia

_ therefore goes into synonymy with Microrutilia.

Townsend’s generic name Prosenostoma is based upon a misidentification of
Brauer & Bergenstamm’s species Senostoma flavipes, and Townsend’s (1932 : 39;
1938 : 420) flavipes (described in the Manual of Myiology as having haired para-

_ facials) is the species hirticeps Malloch. The true flavipes Brauer & Bergenstamm

has bare parafacials, and is a quite different species. However, both Brauer &

| Bergenstamm’s true flavipes and Townsend’s misidentified flavipes (=hirticeps) are

here treated as consubgeneric, and no nomenclatural difficulty arises from Town-

,_ send’s misuse of the name flavipes: Prosenostoma enters into new synonymy with

Microrutilia.

88 R. W. CROSSKEY

Townsend’s error arose from mis-recognition of the types of flavipes. Senostoma
flavipes was described by Brauer & Bergenstamm (1889 : 126) from a gf anda &
specimen from ‘Neu Holland’ and Engel (1925 : 375) recorded these specimens as
‘Typen No. 38 u. 39’. At the same time Engel noted that the Vienna Museum
contained two @ specimens collected by Thorey in Western Australia in 1864; it was
these specimens that Townsend wrongly considered to be the types of flavipes
Brauer & Bergenstamm. In his 1932 work, in which the name Prosenostoma was
proposed, Townsend recorded a ‘Female Ht [holotype] in Wien, labelled ‘““‘Thorey:
1864: Austra. occid.”’ ’ and in the 1938 work (Manual of Myiology, 7 : 420) he again
recorded ‘Ht female from West Australia in Vienna’; Townsend’s notes in the United
States National Museum, Washington, show that there were two specimens, both Q,
with the data ‘Thorey 1864 Austra. occid.’, one being his ‘holotype’ in Vienna and the
other a specimen taken by Townsend from the Vienna collection for his own genotype
collection (the specimen that Townsend obtained from Vienna is still in U.S.N.M. and
has been examined). The two 2 specimens from Western Australia mentioned by
Townsend in his notes (one his supposed holotype) are the same two 9 as Engel had
recorded as ordinary specimens without type-status. The true flavipes types are
the specimens recorded by Engel as “‘Typen’ from “Neu Holland’; these have the
parafacials bare and are a different species from the ‘Thorey 1864 Austra. occid.’
females, apparently being specimens of R. (M.) hirticebs Malloch. A lectotype is
designated from the true type-material of flavipes elsewhere in this paper (see
p. 121). Comparison of this specimen with the lectotype of fulviventris Bigot shows
that flavipes must fall as a synonymy of fulviventris.

Brauer & Bergenstamm (1889) completely misunderstood Macquart’s genus
Senostoma (type-species S. variegata Macquart), which is not even a Rutiliine
(Paramonov, 1968 : 384; Crosskey, 1971 : 291), and were seriously in error to place
flavipes in this genus. In reality it belongs to the genus-group segregate later
described by Townsend as Microrutia. Unfortunately both Engel (1925 : 374)
and Malloch (1929 : 305, 1930 : 109) followed Brauer & Bergenstamm’s erroneous
interpretation of Senostoma, and applied the name Senostoma to the concept which
should correctly be called Microrutilia; hence the entry of Senostoma of authors, not
Macquart, in the foregoing synonymy of Microrutilia.

Microrutilia is an easily recognized subgenus because of the combination of small
size, three postalar setae, conspicuously haired arista, very prominent nasute
epistome and bulbous facial carina, and in the male the unusually prominent sternite
5 and hypopygium. It appears to be most closely related to Grapholostylum with
which it shares a large number of characters (compare diagnoses), and it is possible
that new species discovered in the future may show intermediate characters making
it necessary to amalgamate the two subgenera. At present Microrutilia is easily
distinguished from Grapholostylum by the differently formed ¢ sternite 5 (Text-fig.
32) and by the short membranous distal part of the aedeagus (which is shorter than
the sclerotized proximal part); other differences include the lack of white spotting
on thorax, the presence normally of two posterior intra-alar setae (though there is
variability and some specimens of Microrutilia have only a single fost 7a seta as in
Grapholostylum), and the more strongly developed proclinate orbital bristling (nor-

}

REVISIONARY CLASSIFICATION OF RUTILIINI 89

mally two pairs in Microrutilia females and one or none in Grapholostyium) and
sternopleural bristling (commonly 2 + 1 sternopleurals in Microrutilia and almost
always only 1 + 1 in Grapholostylum).

An interesting feature of the chaetotaxy of Microrutilia (apart from the unusually
long and strong bristles) is the constancy of the posterior dorsocentral setae. In
most Rutilia s.1. the number of fost dc is rather variable within species, and therefore
within any particular subgenus, and the degree of development may differ between

| sexes or on one side of the scutum from the other in the same specimen, but in

Microrutilia there are very constantly four long strong post dc setae regularly spaced
(as they would be in a higher Tachinid with this number of post dc setae) ; the same

_ constancy of four unusually long strong post dc setae (combined with four rather
| strongly developed humeral setae) occurs in Grapholostylum, and perhaps is another

indicator of close phyletic relationship between the two subgenera in spite of the
rather different male sexual characters of Microrutilia and Grapholostylum.

Some species of Microrutilia have a superficial resemblance to some Chrysorutilia
species because of their metallic green upper occiput and postbuccae, and because

_ of the very convex non-sulcate end of the abdomen (T5); but the presence of three
| postalar setae (instead of 4 or 5 as in Chrysorutilia), the lack of pteropleural hair

anterior to the post stpl seta, and the short sparse hairing of the suprasquamal ridge

| (contrasted with the long dense bushy hairing of Chrysorutilia) readily distinguishes

Microrutilia from Chrysorutilia. In Microrutilia, also, there are strong erect setae
present among the hair of abdominal T5, whereas in almost all Chrysorutilia species
this tergite bears fine hair only. The shape of the ¢ sternite 5 is also different in the
two subgenera.

Correct association of the sexes in Microrutilia is especially difficult, particularly

| as the males and females of many if not most of the species (Hirticeps is an exception)
_ appear to be sexually dimorphic in leg colour; females have the legs reddish yellow,
| but males have the legs partly or completely darkened (mainly black or brownish
_ black at least on the coxae, parts of the femora and the tarsi). Some of the nominal
| species in the subgenus are based on ¢ primary types and others on 9 types, and it is
| well-nigh impossible at present (in the absence of bred material or good series

collected in the same place at the same time) to be sure how the females correlate

| with the males; it is almost certain, though, that some of the names involved are
_ synonyms of each other. Some authors have already established synonymies based

on guess-work correlations: Brauer (1899 : 513) placed flavipes as a synonym of

| minor; Austen (1907 : 345) placed liris as a synonym of minor; Engel (1925 : 374)

‘placed flavipes as a synonym of ruficornis; and Townsend (1938) accepted Austen’s
and Engel’s synonymies. During the present work it has been found that there is a

_ very difficult complex of species involved that are all very closely alike, though

differing on male genitalia, and that it is almost impossible to say which females
associate with the different species recognizable on male genitalia. Although at
least one of the previously established synonymies is almost certainly correct, viz.

_ that of liris with minor, it seems best to regard all the names based upon female

types as valid for distinct species until such time as really good evidence is available

for positive association of males and females; in the absence of such evidence I here

go R. W. CROSSKEY

cite the relevant names as valid in the list of included species and in the key to
species.

An interesting feature of most Microrutilia species is the relatively large size of the
epandrium of the g hypopygium and its exceptionally rotund form; this is well seen,
for example, in Text-fig. 62, showing the apical view of the epandrium of R. minor.
The large epandrium tends to make the genitalia appear very prominent when in
situe, an appearance enhanced by the unusually prominent ¢ sternite 5.

INCLUDED SPECIES

Rutilia (Microrutilia) cupreiventris Malloch stat.n. AUSTRALIA (New South
Wales). [Holotype examined].
R. (M.) fulviventris Bigot. AusTRALIA (‘New Holland’, Tasmania). [Lecto-
type examined].
flavipes (Brauer & Bergenstamm) syn.n. [Lectotype examined].
R. (M.) hirticeps Malloch. AustraLiaA (New South Wales, Victoria, Western
Australia). [Holotype examined].
pallens Curran syn. n. [Holotype examined].
[flavipes Brauer & Bergenstamm sensu Townsend. Misidentification.]
R. (M.) liris (Walker). AusTRALIA (Tasmania). [Holotype examined].
R. (M.) media Macquart. AUSTRALIA (Tasmania to New South Wales). [Lecto-
type examined].
vuficornis (Macquart) syn. n. [Holotype examined].
R. (M.) minor Macquart. AUSTRALIA (Tasmania). [Lectotype examined].
R. (M.) nigriceps Malloch. AustTRALiaA (New South Wales). [Holotype exa-
mined].
R. (M.) nigripes (Enderlein) comb. n. Austraia (A.C.T., QUEENSLAND).
[Lectotype examined].

KEY TO SPECIES OF THE SUBGENUS MICRORUTILIA

[Note: Only a small number of female specimens has been available, and some of the key
characters given for females may not be found to be constant with longer series.]

1 Parafacials completely haired. ¢ surstyli and cerci as in ae, 63. Legs of both

sexes entirely reddish yellow . ; : R. hirticeps
— Parafacials bare. ¢ surstyli and cerci differently shaped. Legs sere blackish

brown in g, reddish yellow with dark tarsi in 9. : : 2
2 Femora reddish yellow (9) : : : : é 3
— Femora blackish brown, at most a little reddish basally (33 ) 6

3 Abdomen bicolorous, dark green to greenish black dorsally and elon prange on n most
of venter; last abdominal tergite (T5) covered with thick whitish pollinosity and
therefore contrasting in colour with remainder of abdominal dorsum. Pleural
regions reddish yellow with only slight golden green and violaceous metallic glints.
Mesopleural hair pale yellow (except for a few dark hairs anterodorsally)
R. fulviventris
— Abdomen unicolorous, golden green or emerald or cupreous green on both dorsum and
venter; last visible tergite not noticeably pollinose, therefore shining like remainder
of abdomen. Pleural regions mainly metallic golden green or blue-green. Hair
of mesopleuron black (except for some pale hair along extreme anterior edge) : 4
4 Basicosta bright orange. Parafrontals not metallic at upper ends. Sternopleural

REVISIONARY CLASSIFICATION OF RUTILIINI gt

hair pale yellow. Abdomen partly cupreous. Proclinate orbital setae in one pair
only or none definitely developed. 1 + 1 sternopleural setae
[Possibly the 2 of R. media] R. cupreiventris

Basicosta blackish brown or at least dark brown anteriorly. Parafrontals showing
metallic green colour at upper ends on to vertex. Sternopleuron with hair entirely
black or black on upper part around and between the stp/ setae. Abdomen all
golden green oremerald. Two pairs of strongly developed proclinate orbital setae.
Normally 2 + 1 stpl setae (occasionally only 1 + 1) .

Abdominal tergites 3 and 4 without discal setae and with entirely recumbent hair:
T3 without median marginal setae Sternopleural hair all black Ground colour
of lower parts of parafrontals reddish and not noticeably contrasting in any light
with the reddish yellow ground colour of parafacials. Setae of the upper part
of the postocular row short and straight . : : . : Undetermined sp.

[Possibly the 2 of R. nigripes]

Abdominal tergites 3 and 4 with some long fine irregular discal setae distinctly
developed among the hair, the hair sub-erect medially on T3 and erect all over
T4; T3 with a row of about four long fine median marginal setae. Sternopleural
hair black around and between the stp setae but pale yellowish on mid part of
sternopleuron. Ground colour of parafrontals blackish and in some lights con-
trasting rather abruptly with reddish yellow ground colour of parafacials. Setae
of the upper part of the postocular row rather amie and fine and curved slightly
forwards. : é ; : } : E R. liris

[Probably the ? of Vee ‘nines

Hair of pleural regions and abdominal venter pale yellow. Hair of fore coxae

mainly golden : : Undetermined sp. or spp.
[Possibly g of R. fulviventris. 3 genitalia ‘souilar to R. nigripes. Specimens
from Queensland have extremely narrow upper frons as in nigripes, measuring
0:05-0:06 of head-width. Other specimens from Victoria have frons at narrowest
from 0-065-0-080fhead-width. Queenslandspecimens belong to R.vuficornis sensu
Enderlein, not Macquart. They may possibly be a colour polymorph of nigripes.]

Hair of pleural regions and abdominal venter almost all black. Hair of fore coxae
black (at most a few golden hairs at extreme base and apex) ;

Ground colour of parafrontals entirely blackish. Very small species (length es mm)
with exceptionally long fine bristling and oe 6 surstyli and cerci as in ae
62 or Text-fig. 109 : 8

Ground colour of parafrontals tawny reddish or brownish, at most only a little
blackish at upper ends. Larger species (length 9:5-15 mm) with vestiture not
exceptionally long and fine. ¢ surstyli and cerci of different shape (Text-figs
60 & 61) a ‘ ‘ ‘ 9

Ground colour of parafacials blackish ‘and concolorous with ey 6 surstyli
and cerci as in Text-fig. 109, surstyli with sharp apical point and cerci sinuous in
profile. rons at narrowest point distinctly wider than third antennal segment

R. nigriceps

Ground colour of parafacials yellow and rather sharply contrasting with the blackish
parafrontals. 4 surstyli and cerci as in Text-fig. 62, surstyli rounded apically and
cerci straight in profile. Frons at narrowest point ee lees in width to or very
slightly narrower than third antennal segment . : : . R. minor

Eyes exceptionally strongly approximated, interfrontal area almost completely
obliterated at its upper end, frons at narrowest point only 0:05-0:06 of head-width.

6 genitalia with free apical parts of cerci not much longer than the contiguous
basal parts, and the outer margins of the bases of the cerci rounded (Text-fig. 61).
[Queensland] i : ; R. nigripes

Eyes less strongly approximated, upper part of interfrontal area narrow but distinct,

frons at narrowest point 0-075-0-08 of head-width. ¢ genitalia with exceptionally

nn

“I

92 R. W. CROSSKEY

large heavy cerci (Text-fig. 60) in which the free apical parts are much longer than
the contiguous basal midline of the cerci; the outer margins of the basal parts of
the cerci rather straight. [Tasmania and New South Wales] . : . R. media

SUBGENERICALLY UNPLACED SPECIES-GROUP Taxa IN RUTILIA S.t.

These are two, micropalpis Malloch, 1929 and scutellata Enderlein, 1936. Each is
briefly discussed below.

Rutilia micropalpis Malloch

This little-known species differs from all other species of Rutilia s.1. so far described
by having the palpi (as the appropriate specific name indicates) exceptionally small;
they are very inconspicuous in the buccal cavity and their length is at most only
about half the normal palpal length in Rwutilia. I would not exclude the species
from any named subgenus on this character alone, but on the basis of all the other
characters shown by micropalpis in combination it is impossible to place the species
at all satisfactorily in any one of the seven subgenera here recognized, and it seems
best to leave micropalpis subgenerically unplaced for the time being. The male is
unknown to me, and so far I have seen only four female specimens (holotype and
paratype in Australian Museum and two specimens in British Museum (Natural
History)). Other than the palpal character these are the features shown by micro-

palpis:

Head pollinose, non-metallic; parafacials bare; 2 with one pair of strong proclinate orbital
setae; arista micropubescent; four humeral] setae (one sometimes very weak); one to three post-
humeral setae (variable each side on same specimen); two or three post ia setae (if three then
anterior one very small); no supernumerary prescutellar setae; four or five postalar setae;
suprasquamal ridge with short black hair, rather sparse; scutellum convex, with four or five
paiis of marginal setae and with well developed preapicals; pteropleuron not haired in advance
of hind stpl seta; two sternopleural setae (1 + 1, anterior one exceptionally strong); prosternal
membrane and anterior edge of prosternum with long pale hair; hind tibia with short very
regular and close-set ad fringe along its basal three-fifths and with or without one strong ad
seta inserted at the end of the fringe, with three or fom strong pd setae; abdominal T3 with
one pair of median marginal setae; T5 without depression and with a median transverse row
of very strong erect setae; body colour dark blackish green with coppery to purplish reflections
(mainly on mesonotum and scutellum) and thin whitish pollinosity over humeral area, meso-
pleuron, and (very thinly) on tergite bases.

If the foregoing list of micropalpis characters is compared with the subgeneric
diagnoses it is evident that the closest fit lies with subgenus Donovanius, though the
species fails to conform with typical species of this subgenus by having short sparse
suprasquamal ridge hairing, some hair present on the prosternum, by the very short
hind tibial fringe and several strong hind tibial fd setae, and by the lack of a depres-
sion in the last abdominal tergite. R. micropalpis seems to combine some of the
characters of Donovanius with some of Rutilias.str. There is no suggestion, or very
little, of any affinity with the other subgenera, even though these possess four or
more postalar setae as in micropalpis.

It would not be unjustified on the basis of its unusual palpi and combination of
other characters to place micropalpis in a separate new subgenus, but one is not

REVISIONARY CLASSIFICATION OF RUTILIINI 93

proposed at present because the male is still unknown; the male genitalia might well
give a better clue to the affinities than any of the female characteristics known so
far. Ifit should prove, as seems possible, that the male of micropalpis has genitalia
of the Donovanius type (with large heavy foliaceous surstyli) then it might be best to
assign micropalpis to this subgenus in spite of its somewhat aberrant features such
as the reduced palpi.

Chrysorutilia media var. scutellata Enderlein

This variety was described from a single female (holotype) collected at Adelaide,
South Australia, and the description consists of the five words ‘Scutellum dunkel
rostfarben ohne Metallglanz’. The holotype is probably in the Berlin Museum
where it should correctly be located, but was not found there when other Enderlein
types were borrowed for this work. In the absence of the type I cannot place the
name scutellata, but it presumably applies to a species of Chrysorutilia (most probably
asasynonym). It almost certainly has nothing to do with media Macquart, which
belongs in the subgenus Microrutilia and was clearly misidentified by Enderlein
(though specimens named as media by Enderlein have not been seen).

Genus AMPHIBOLIA Macquart

Amphibohha Macquart, 1843 : 278 (121). Type-species: Amphibolia valentina Macquart, 1843,
by original designation and monotypy.

Diacnosis. Facial carina widest above middle and distinctly convergent ventrally, rounded
on anterior surface, upper part often rather bulbous. Epistome moderately to strongly promi-
nent, face strongly excavate in profile between epistome and carina. Head of ¢ not holoptic,
upper eye facets not enlarged, separated by a distance much greater than width of facial carina.
Genal dilation well developed. Parafacials haired or bare. Buccal opening normal, very much
wider than facial carina. Head pollinose, non-metallic. Arista pubescent. Palpi normal,
fully developed. Mentum short and rather broad in profile with subparallel sides. Prosternum
bare, prosternal membrane bare or haired. Scutellum with apical pair of setae inserted at
lower level than other marginal setae (very rarely absent); total of 4-6 (7) pairs of marginals;
disc of scutellum not flattened. Postalar callus with 3-4 setae (occasional specimen with small
supernumeraries in addition). Postalar wall bare. Suprasquamal ridge thickly haired. Upper
calypter normal. Tegula with normal long wiry posterior setulae. Costal base not explanate.
Abdomen with marginal vestiture of tergite venters weak and semi-recumbent (directed back-
wards), if slightly spiniform (as in assimilis) then not directed vertically downwards; T3 with
median marginal setae (often numerous and in strongly developed transverse row) ; intermediate
tergites with discal setae (absent in occasional specimens). T5 convex above and broadly trun-
cate subconical, without depression or at most with only very slight apicomedian hollowing.

DISTRIBUTION. Australia and Tasmania, Lord Howe Island, New Guinea.
Occurring in Australia from Western Australia to Victoria and New South Wales;
Tepresented, but apparently poorly, in Queensland.

Discussion. It is by no means certain that Amphibolia ought to be considered
generically distinct from Rutilia s.1., and it is maintained here as a separate genus
with considerable doubts as to whether this is fully justified. In the past none of the
earlier workers on the Rutiliini have doubted its generic distinctness, but the fact is

94 KW, GCROSSIEGE ¥

that most of the characteristics of Amphibolia conform exactly or very closely with
those of Rutilia in the wide sense and that it is difficult to find really convincing
characters for generic separation. The type-species, A. valentina, remained for
many years the only known species and seemed very distinct from all the Rutilia
species because of the exceptionally unusual and conspicuous bold black-and-white
pattern of the thorax and abdomen, a pattern that elsewhere in the Rutiliinioccurred
only in the superficially similar species Formosia speciosa Erichson; now, however,
that more species are known through the work of Paramonov (1950, 1968) it is clear
that valentina is not nearly so distinctive as it seemed, and that other species of
Amphibolia such as A. campbelli Paramonov, though having the black-and-white
pattern, have features of the facies, strength and arrangement of the chaetotaxy
and so forth that suggest rather close affinity to, particularly, the segregates Param-
phlibolia and Chaetogastrina. The two species contained in these hitherto monotypic
genera look very different from Amp/ibolia when arranged in a collection, because
they lack the black-and-white pattern, yet they do not differ (apart from pattern)
from typical Amphibolia on their structural characters to any greater extent than do
many of the species of Rutilia or Formosia differ from their more typical congeners.
Considering the range of known species in Amphibolia, Paramphibolia and Chaeto-
gastrina as a whole, therefore, it seems best to treat them in an equivalent manner to
the species here placed in Rutilia s.1. by merging them into a single genus and widen-
ing the generic definition of Amp/ibolia accordingly. But the form of the male
hypopygium and fifth sternite which is essentially very similar in assimilis and
stolida (type-species respectively of Paramphibolia and Chaetogastrina) differs
slightly from that of typical Amphibolia species, and for this reason (taken in
conjunction with the striking pattern difference) it is considered best to recognize
two subgenera, distinguished by the same order of difference as that distinguishing
the various subgenera here recognized in Formosia and Rutilia.

The main characteristics distinguishing Amphibolia s.l. from other Rutiliine
genera are as follows: suprasquamal ridge haired (distinction from Formosia,
Formodexia, Chetogaster, Rutilodexia and Prodiaphania), postalar callus with three
or four setae (distinction from Chetogaster), palpi normal and arista micropubescent
(distinction from Prodiaphania); head entirely pollinose, non-metallic (distinction
from Chrysopasta) ; facial carina contracted or evanescent ventrally (distinction from
Rutilia s.1.); intermediate abdominal tergites (T3 and T4) almost always with discal
setae (distinction from all other genera except Chetogaster).

Amphibolia s.l. is most nearly allied to Rutilia, and the distinctions between these
time-honoured genera are rather intangible. In general in Rutilia the facial carina is
rather broad, often flattened on the fore surface, and not strikingly evanescent at its
ventral end, whereas in Amphibolia the carina is conspicuously best developed on the
upper part (which may be obviously bulbous) and weakly developed with the sides
much convergent at the ventral end. Nearly always in Amphibolia there are a few
erect discal setae present on tergites 3 and 4, but occasionally specimens lack
abdominal discal setae on these tergites or one of the tergites may have only a
single seta; on the other hand discal setae are apparently never present on the
intermediate tergites in Rutilia (though doubtless some specimens of this genus will

REVISIONARY CLASSIFICATION OF RUTILIINI 95

ultimately be found that possess adventitious discals). But presence or absence of
discals on T3 and T4 is a useful rule-of-thumb distinction between Amphibolia and
Rutilia. Those Rutilia (a few species) which have the suprasquamal ridge bare are
immediately separable from Amphibolia by this character (all known species of
Amphbolia have the suprasquamal ridge fully haired). Likewise those Rutilia
species that have a very broad medially depressed T5 are at once separable from
Amphibolia, which does not have a deep median depression in this tergite (the last
tergite in all Amphibolia species is truncate subconical with scarcely any trace of
flattening or hollowing medially at the tip).
The two subgenera of Amphibolia are distinguished by the following key:

KEY TO THE SUBGENERA OF AMPHIBOLIA

t Thorax and abdomen with bold black-and-white pattern. Lobes of fifth sternite of
3 simple. ¢ hypopygium with surstyli shorter than cerci (Text-figs 87-89).
Apical pair of scutellar setae odes as strong as other scutellar marginal
setae . . AMPHIBOLIA Macquart (p. 95)
- Thorax and abdomen without such pattern. Lobes of fifth sternite of g with small
tooth or prong on inner edge near apex (Text-figs 34 & 35). dg hypopygium with
surstyli longer than cerci (Text-figs 85 & 86). Apical pair of scutellar setae much
weaker than other marginal setae (sometimes absent)
PARAMPHIBOLIA Brauer & Bergenstamm (p. 100)

Subgenus AMPHIBOLIA Macquart

Amphibolia Macquart, 1843 : 278 (121). Type-species: Amphibolia valentina Macquart, 1843
by original designation and monotypy.

Diacnosis. Thorax black with bold discrete white-pollinose spots on dorsum. Calyptrae
_ black or brownish black. Scutellum with apical pair of setae subequal in size to other marginal
setae. Abdomen conspicuously patterned in contrasting black and white-pollinose araes.
Thoracic and abdominal chaetotaxy not spiniform. Lobes of g fifth sternite simple, without
subapical tooth oninneredge. ¢g hypopygium with short broad surstyli (often slightly bifurcate)
which are shorter than cerci.

DISTRIBUTION. Australia and Tasmania, Lord Howe Island, New Guinea.

_ Discussion. This subgenus includes some of the most attractive and beautifully

marked flies among the whole of the Rutiliini which are at once recognized by eye
_ from their bold black-and-white coloration of the thorax and abdomen combined
with their bright yellow heads. Not all species, however, which belong in the
_ subgenus are quite so distinctive, for some have the white areas less boldly displayed
than others, and not all of them have the yellow head colour. In general the species
fall into two groups, one including the type-species (valentina) in which the head is
mainly yellow or at most reddish brown and the abdomen has (as part of the pattern)
discrete rounded black spots, and another including species in which the head
ground colour is distinctly black and in which the abdomen (though possessing a
' pattern) lacks definite isolated black spots; these two groups, which are both Austra-
lian, are however interconnected by a new species (papuana) here described from

New Guinea, which has the black ground colour of the head but also has discrete

96 Ro Wi.) CROSSIGCE

black abdominal spotting. As there is intergradation between the species it is not
proposed to recognize formal species-groups.

The species of Amp/ibolia s. str. show some variability in the development of the
chaetotaxy. In A. ignorata the bristling is especially weak and exceptionally short
on the head and abdomen, and the head bristling is also very weak in A. valentina
(in both these species the frontal setae are so short and fine that the rows are not
cruciate and scarcely meet at the tips), but in A. campbelli all the setae are long and
strong and the chaetotaxy of this species is almost identical with that of A. stolida in
the subgenus Pavamphibolia. In A. campbell: (and also the closely similar A. wilsont)
the hairing as well as the chaetotaxy is more strongly developed than in other
species, the parafacials being entirely haired (these are bare in all the species in which
the head ground colour is either all bright yellow or all blackish). The pollinosity of
the upper parafacials and lower parafrontals in A. campbelli has a shifting appearance
according to the direction of the light which is unusual in Rutiliini, but recalls a
rather similar condition found on the head of Chrysopasta elegans, and it is possible
that the monotypic genus Chrysopasta has closer affinity with Amp/bolia than with
any other genus of Rutiliines.

An astonishing convergent resemblance exists between the black-and-white boldly
patterned Amphibolia s.str. species and some other Calyptrate flies with similar
patterning. Inthe subgenus Euampiibolia Townsend (q.v.), a segregate of Formosia
s.l., and in the Ameniine Calliphorid genus Formosiomima Enderlein (see Crosskey,
1965), the thorax and abdomen has a black-and-white pattern formed in exactly
the same way as in Amphibolia. In all of these flies the thoracic dorsum is black
with thickly pollinose white marks in pairs on (1) the notopleural to humeral area,
(2) the prescutum, (3) the supra-alar area, and (4) on the scutum submedially (these
last sometimes evanescent), to which basic pattern there may be superimposed
additional white-pollinose marks (usually in the form of one or two longitudinal
vittae between the paired prescutal sublateral marks). The abdominal patterns
are less constant throughout the range of species, though constant within a species,
but the nature of the pattern is the same—being formed of areas of extensive thick
overlay of white pollinosity contrasting with areas of black ground colour devoid of
such pollinosity. In all three taxa of these apparent mimics, Amphibolia, Euamphi-
bolia and Formosiomima, part of the abdomen appears black-spotted, each black
spot in reality being an island (or a confluent pair of islands) not covered by the
white-pollinose overlay. Some specimens of all three taxa also have a suggestion of
greenish or violaceous tinge showing through the pale pollinose areas (an interference
effect of the pollinosity overlying the blackish ground colour).

Paramonov (1968 : 357) states that Amp/ibolia species are parasitic on larval
cockchafers. This is most probably correct, though Paramonov’s evidence came
only from a specimen of A. valentina in the CSIRO collection, Canberra, that was
collected in Victoria and bears a label reading ‘larvae of these flies parasitic on
cockchafer grubs’.

INCLUDED SPECIES
Amphibolia (Amphibolia) albocincta (Malloch). AusTRALIA (Australian

REVISIONARY CLASSIFICATION OF RUTILIINI 97

Capital Territory, New South Wales). [Holotype examined].

A. (A.) campbelli Paramonov. AvusTRALIA (Australian Capital Territory, New
South Wales, Victoria). [Holotype examined].

A. (A.) commoni Paramonov. AUSTRALIA (New South Wales, Victoria).

A. (A.) ignorata Paramonov. AUSTRALIA (Western Australia to Victoria to
Queensland); Lorp Howe Isianp. [Holotype examined].

A. (A.) papuana Crosskey sp.n. New Guinea. [Described below].

A. (A.) valentina Macquart. AvusTRALIA (Western Australia to Victoria to
Queensland, Tasmania).

vidua (Guérin-Méneville). [Syntypes lost].
A. (A.) wilsoni Paramonoy. AUSTRALIA (Victoria).

KeEy To SPECIES OF THE SUBGENUS AMPHIBOLIA

Parafacials bare

Parafacials completely haired P 6
Head yellow to orange-red with thick yellow to golden orange pollinosity Hair of

genae and postbuccae golden yellow : 3
Head black (at most slightly reddish on genae) with whitish c or ashy grey pollinosity

Hair of genae and postbuccae black. [Only 92 known] . 4

Last visible abdominal tergite (T5) entirely black. Submedian pair of black spots on
T3 standing close together and normally coalesced before hind margin of the
tergite. Pattern of abdominal T4 consisting of two very large semi-circular black
marks which meet and fuse in the mid-line (usually also a pair of very minute incon-
spicuous black dots in the white-pollinose area near the mid fore margin of the
tergite). Vertex of § o-11-0-12 of head-width. Surstylus of ¢ genitalia as in
Text-fig. 87 . A, ignorata
Last visible ppc tergite (Ts) with black sindi-igiht pattern, sides of the tergite
thickly white pollinose to a variable extent. Submedian pair of black spots on T3
not meeting each other at hind margin of tergite, very clearly separated by white
pollinose area which extends back in mid line to hind margin of tergite. Pattern
of T4 consisting of four large black spots standing against hind margin of tergite, an
isolated lateral pair and an inner pair of rounded spots which fuse in the mid-line
(occasional specimen may have lateral spot larger than usual and just meeting with
the inner spot); T4 in addition always with a pair of small but distinct submedian
black dots near fore margin. Vertex of f wider, 0:14-0:15 of head-width. Sur-
stylus of $ genitalia as in Text-fig. 88 ‘ A. valentina
Abdominal T3 with a thick covering of whitish pollinosity and a black spot pattern
consisting of a large transverse median spot and a pair of lateral spots standing on
the hind margin of the tergite (in addition a very small black V-shaped area
encroaching on white pollinose area anteriorly in the mid-line). [New Guinea]
A, papuana sp. n. (p. 98)
Abdominal T3 without a pattern of black spots, the black areas of the tergite in
form of transverse bands. [Australia] . 5
Abdominal T3 almost completely covered with thick, white pollnilasite foal oe
narrowly black against hind margin), abdomen therefore appearing to have a

broad white fascia sub-basally : ? A. albocincta
Abdominal T3 mainly black, with only a narrow : basal band of whitish pollinosity
which is discontinuous at the mid-line . , ; A. commoni

Vertex of J o-1 3-0'14 of head-width. Prescutum aed a mean whitish pollinose vitta
which (seen in some lights) is more or less continuous to the transverse suture.
Surstyli and cerci of f genitalia as in Text-fig. 89 . - : : A, campbelli

98 R. W. CROSSKEY

— Vertex of § unusually narrow, about 0-09 of head-width. Median whitish vitta of

the prescutum evanescent posteriorly, not reaching to suture . : . A. wilsoni
[This species is known only from the g holotype. The genitalia have not been
examined. ]

Amphibolia (Amphibolia) papuana sp. n.

@. Head. Ground colour mainly blackish, except for yellowish facial carina and reddish
brown genal and postbuccal regions; parafiontals and interfrontal area black; head pollinosity
white, thinner on parafrontals than elsewhere (in some lights lower end of parafrontal shows
distinct line of demarcation where thicker pollinosity on parafacials and bottom end of para-
frontal meets area of thinner pollinosity on parafrontal). Parafrontal and occipital hair black,
genal and post buccal hair dark reddish brown. Frontal setae short and very fine, tips of oppo-
site rows not meeting; one pair of divaricate ocellar setae; two pairs of well developed proclinate
orbital setae. Vertex 0-24 of head width. Facial carina longer than epistome, narrow and
nearly subfusiform but slightly widened at level with junction of second and third antennal
segments and very slender on ventral third; facial profile moderately excavate between epistome
and carina. Gena exceptionally broad, about 0-55 of eye-height. Parafacials very broad,
about three and a half times wider than third antennal segment, entirely bare. Antennae
slightly elongate, falling short of mouth-margin by about two-thirds of length of third segment;
basal segments reddish with some black infuscation on inner edge of second segment, third
segment blackish brown except for some bright orange colour at base; arista with short pubes-
cence. Palpi tawny yellow. Thorax. Brownish black to black with bold white spots con-
spicuous to naked eye. Spots arranged as pair of large humeral-notopleural marks, pair of
sublateral prescutal vittae (in which each vitta strongly contracted medially so that to naked
eye the white vitta appears as separated spots), pair of slender median white vittae on prescutum
between the sublateral vittae, pair of supra-alar spots on scutum and pair of submedian spots
on scutum (these less boldly white than other spots); in addition prescutum with pair of very
small spots standing against transverse suture just mesad of humeral-notopleural spots; meso-
pleuron and sternopleuiron with large white spot; seen from in front prescutum shows trace of
blue-green colouring under the anterior parts of the sublateral white vittae, and seen laterally
scutum shows trace of coppery violaceous glints around supra-alar spots. All thoracic hair
black. Chaetotaxy rather weak, three pairs of presutural acrostichal setae differentiated
(standing on median white lines), two post ia setae, I + 1 sternopleural setae, no setae developed
on inner half of humeral callus (only two setae on outer half). Scutellum with very strong
crossed apical setae, three or four pairs of other marginals, and with well developed preapicals.
Wings. Basicosta dark. Wing membrane hyaline except for boldly marked basal dark spot
in which cells completely blackened. Calyptrae blackish brown with dark brown marginal
hair. Legs. Blackish brown with all black hair. Chaetotaxy weak, most setae short and
fine; mid tibia with only one distinct ad seta; hind tibia with very short weak ad fringe and with
two main ad setae and two pd setae. Abdomen. With black-and-white pattern. Tr + 2
and T5 entirely black. T3 with thick covering of whitish pollinosity over most of surface,
except for very small median anterior V-shaped black mark and three black spots against hind
margin; black spots arranged as lateral pair on extreme sides of abdomen and a broad median
spot. T4 black with a transverse fascia of thick whitish pollinose covering on anterior quarter
or third, the pale pollinose area extending back to occupy nearly half the tergite length mid
dorsally and extending black medioventrally to reach the hind margin. Greenish tinge visible
on pale pollinose areas of T4 dorsum, and venter distinctly green under the pale pollinosity;
black areas of venter with violaceous tinge in some lights. Pale pollinose parts of T3 and T4
together giving abdomen appearance of having a broad transverse pale band contrasting with
the black base and apex. Abdominal hairing black, mainly short, fine and semi-recumbent,
but longer and erect on T5 and medially on T4. T3 with one pair of erect median marginal
setae. T3 and T4 without discal setae, but holotype (only available specimen) has one large
pore (seta itself missing) on the anteromedian V-shaped black mark of T3 near the fore margin

REVISIONARY CLASSIFICATION OF RUTILIINI 99

of the tergite. T5 without trace of median depression, with a transverse row of strong erect
discal setae standing near the centre of the tergite. Measurements. Body length 14:7 mm,
wing length 12-6 mm [holotype only].

3d. Unknown.

MATERIAL EXAMINED

Holotype 9, NEw GuINnEA: Murmur Pass, 8600 ft, x.1961 (W. W. Brandt). In
British Museum (Natural History), London.

DIsTRIBUTION. Known only from the holotype from highland New Guinea.

AFFINITIES. This is the first species of Amphibolia to be discovered in New
Guinea. Although only a single specimen is so far known it has been considered
desirable to describe the new species because its provenance is unexpected and
because it shows very distinctive features which make it intermediate between two
groupings of Amphibolia s.str. species present in the mainland Australian fauna.
Paramonov (1968 : 356) found that the Australian species fell into two moderately
distinct groups, one with mainly yellow head and with black abdominal spots and
the other with blackish head and no black abdominal spots, and he used this distinc-
tion for the first cut in his key to the Australian species. The new species A. (A.)
_ papuana here described has the black head colour of the second of Paramonov’s
groups but has bold black abdominal spots on T3 like the first group, and seems on
this account to occupy a rather intermediate position; on total facies, however, it
appears most closely allied to A. (A.) albocincta (Malloch), a species with dark head
and no black spots from New South Wales. From a zoogeographical viewpoint this
possibility is of interest because it provides another instance of a curious fact of
Tachinid distribution that is becoming more and more evident—that there are
elements in the Tachinid fauna of the New Guinea highlands that are quite disjunct
from the rest of the Papuan fauna but are beyond doubt extremely closely allied to
similar or almost identical forms in the mountainous parts of New South Wales
(with, apparently, a very wide gap in distribution in Queensland).

The holotype of papuana sp. n. differs from typical Amphibolia specimens by
seeming to lack definite discal setae on the intermediate abdominal tergites, but
careful examination shows the presence of a large pore anteromedially on T3,
confirming that abdominal discals can be present in this species. The probability
is that specimens are variable in the development of discal setae in papwana, as in
other species of Amphibolia, and it is to be expected that specimens obtained in
future will not necessarily conform completely with the holotype in this detail of the
_ abdominal chaetotaxy.

_ Asa whole the strength and development of the chaetotaxy in papuana sp.n. is
closely similar to that of A. (A.) ignorata Paramonov, in both species the abdominal
_ and leg chaetotaxy in particular being very weak. It is of interest to note also the
_ close resemblance in the distribution of the pale pollinose overlay of the abdomen in
_ these two species; the abdominal pattern is essentially extremely similar in papuana
and ignorata, differing only in that ignorata has a pair of minute median black spots
' in addition in the centre of the pale pollinose fascia of T4, and the median black spot
of T3 in ignorata is clearly composed of two partially fused spots. But ignorata has
the head bright yellow and the antennae orange and on this feature alone is at once

100 R. W. CROSSKEY

separable from papuana (a specimen in the BMNH collection from Queensland stand-
ing under the name ignorata and having the bright yellow head has the abdominal
pattern exactly as in papuana, and might perhaps belong to a different species: the
specimen is of interest in showing that flies with an abdominal pattern that is an
exact match with that of the New Guinea species occur in mainland Australia, even
though quite different on other characteristics).

Subgenus PARAMPHIBOLIA Brauer & Bergenstamm stat. n.

Paramphiboia Brauer & Bergenstamm, 1891 : 389 (85). Type-species: Rutilia assimilis

Macquart, 1851, by monotypy.

Chaetogastrina Malloch, 1929 : 313. Type-species: Chaetogastrina stolida Malloch, 1929, by

original designation. Syn. n.

Diacnosis. Thorax without bold white pollen spots. Calyptrae semi-translucent yellowish
or brownish. Scutellum with apical setae weak, much smaller than other marginal setae and
sometimes not developed at all. Abdomen not patterned in contrasting black and white-
pollinose areas. Abdominal chaetotaxy strong to slightly spiniform. Lobes of ¢ fifth sternite
with distinct tooth or small sharp prong subapically on inner edge. g hypopygium with sur-
styli as long as the cerci.

DISTRIBUTION. South-eastern Australia and Tasmania only.

Discussion. This subgenus contains only two known species, Amphibolia
(Paramphibolia) assimilis and A. (P.) stolida (Malloch), which are the type-species
of Paramphibolia and Chaetogastrina respectively. Malloch’s description of Chaeto-
gastrina consisted only of the sentence “This genus is very similar to Paramphibolia,
agreeing with it in structure of the head and thorax, but there are no parafacial
hairs below level of bases of antennae’, from which it is evident that Malloch’s only
criterion for generic separation lay in the bare or haired parafacials. There is no
doubt that in the Rutiliini as a whole there are many very closely allied species which
differ in the degree of hairiness of the parafacials, some species belonging obviously
to the same distinctive group having them bare, others partially haired, and others
completely haired. In the present work no subgeneric or generic significance is
attached to bareness or hairiness of the parafacials, a character considered to be
specific only (though a few subgeneric segregates may have the same condition in all
or most of the included species). Since there are no other notable differences between
assimilis and stolida (though their superficial appearance is different because of
different body and hair colouring and because of the different degree of development
of the abdominal setae) these two species are here treated as unquestionably con-
generic and consubgeneric; hence the name Chaetogastrina is placed as a new
synonym of Paramphibolia.

The new synonymy of Chaetogastrina with Paramphbolia is supported by evidence
from the fifth sternite and hypopygium of the male (which were not examined by
Malloch). In both stolida and assimilis each lobe of the fifth sternite is produced
on the subapical part of the inner margin into a small tooth, a development that is
found (to the best of my knowledge) nowhere else in the Rutiliini. In fact through-
out most of the Proseninae the lobes of the male fifth sternite are simple rounded struc-
tures without any specialized developments. There is no doubt that the small

REVISIONARY CLASSIFICATION OF RUTILIINI IOI

blunt sternite tooth in stolida (Text-fig. 34) and the rather more prominent, sharper,
tooth in assimilis (Text-fig. 35) are homologous, and good indicators of a recent
_ common ancestry for these two species. The hypopygium itself is also similar in
the two species in the degree of development of the surstyli: although these are
_ different in shape, and quite conspicuously so, they are large and long in both species
_ and in profile their apices reach to a level with the apices of the cerci. In Amphibolia
_s.str. the fifth sternite lobes are simple and rounded as in other Rutiliini without
_ trace of a tooth, and the surstyli are much shorter than the cerci; hence the genital
structures provide the most positive morphological features for the differentiation of
| Paramphibolia (syn. Chaetogastrina) and Amphibolia s.str.
Paramonov (1954 : 275, 1968 : 367) assigned Malloch’s Chaetogastrina stolida to
| the genus Chetogastey Macquart, and therefore sank the name Chaetogastrina into
_ synonymy with Chetogaster, but he gave no arguments in support of this curious
, conclusion. There is a slight superficial resemblance between stolida and, for exam-
| ple, Chetogaster oblonga (Macquart), but otherwise there is almost nothing to suggest
_ any close relationship between stolida and Chetogaster; stolida has supernumerary
_ setae on the postalar callus, haired suprasquamal ridge, fully haired barette, and
_ other features completely characteristic of the Amphibolia—Rutilia—Chrysopasta
complex of genera, and it is here concluded that Paramonov’s placement of stolida
(and therefore of Chaetogastrina) is much in error. Malloch (1929) was undoubtedly
_ right to emphasize the close resemblance between stolida and Paramphibolia.
The bristling of the atdomen in A. (P.) assimilis is so characteristic that this
| species can be distinguished at a glance on this feature from all other Rutiliini; the
_ setae are unusually thickened and abundant, there being a transverse row of many
_ erect setae across the whole hind margin of T3, and exceptional strong development
of slightly spiniform discal setae on T3 and T4 (these varying in number and
arrangement but at least two always being present and very obvious on each tergite).
Sometimes the strong discals of one or both intermediate tergites form a large cluster
or an irregular transverse row.

Specimens of both assimilis and stolida are rare in collections, but on available
_ evidence the two species appear to be allopatric: assimilis is found in Tasmania and
Victoria, and stolida in the mountainous parts of New South Wales. Nothing is
_ known of the host-relations.

INCLUDED SPECIES

Amphibolia (Paramphibolia) assimilis (Macquart) comb. n. AUSTRALIA
(Tasmania, Victoria). [Lectotype examined].

A. (P.) stolida (Malloch) comb. n. AvusTRALIA (New South Wales). [Holotype
examined].

KEry To SPECIES OF THE SUBGENUS PARA MPHIBOLIA

'1 Parafacials haired. Pleural regions of thorax with yellowish white to golden yellow
hair (some dark hair on mid-mesopleuron). Abdomen mainly pale tawny brown
($) to red-brown (2) with distinct median black vitta. Abdominal chaetotaxy
mainly strongly spiniform. Surstyliof¢ genitalia as in Text-fig. 85. [Tasmania &
Victoria] ; : is . F : ‘ : A, assimilis

102 R. W. CROSSKEY

— Parafacials bare. Pleural regions of thorax with black hair. Abdomen very dark
brownish black with indistinct median vitta. Abdominal chaetotaxy not
unusually spiniform. Surstyli of g genitalia as in Text-fig. 86. [New South
Wales] : 5 : : : : : 2 : 3 A. stolida

Genus CHRYSOPASTA Brauer & Bergenstamm

Chrysopasta Brauer & Bergenstamm, 1889 : 152. Type-species: Chrysopasta versicoloy Brauer
& Bergenstamm, 1889 [=Rutilia elegans Macquart, 1846], by original designation and mono-
typy.

Roederia Brauer & Bergenstamm, 1893 : 98. Type-species: Chrysopasta versicoloy Brauer &
Bergenstamm, 1889 [=Rutilia elegans Macquart, 1846], by monotypy (see text for explana-
tion). [Junior homonym preoccupied by Roederia Mik, 1881}.

Echrysopasta Townsend, 1932 : 39. Type-species: Rutilia elegans Macquart, 1846, by original
designation.

Euchrysopasta: Paramonov, 1968 : 372-373. Incorrect subsequent spelling of Echrysopasta
Townsend, without status in nomenclature.

Diacnosis. Facial carina with prominent bulbous upper part contracting ventrally to a
sharp narrow ridge. Epistome very strongly prominent, subnasute, face in profile deeply
concave between epistome and carina. Head of g not holoptic and upper eye facets not en-
larged, frons much wider than facial carina. Parafacials densely long haired, hairing virtually
continuous with that of genae. Buccal opening normal, much wider than facial carina in both
sexes. Genal dilation strongly developed, extending forwards to or just beyond level of front
of eye. Head pollinose anteriorly and on postorbits (pollinosity of lower parafrontals and
postorbits with shifting chequered appearance), metallic on occipital and postbuccal regions.
Arista micropubescent. Palpi medium size, much shorter than the unusually slender proboscis.
Proboscis with slender mentum tapering slightly in profile towards labellae. Prosternum and
prosternal membrane bare or haired (prosternum even strongly setose in some specimens).
Scutellum with apical pair of setae inserted at lower level than other marginal setae; total of
4-5 pairs of marginals; disc of scutellum convex. Postalar callus with three setae. Postalar
wall bare. Suprasquamal ridge haired. Upper calypter normal. Tegula with normal long
wiry posterior setulae. Costal base not explanate. Abdomen with marginal vestiture of
tergite venters hairy only and semi-recumbent, no development of strong setae; T3 without
median marginal setae in gf or at most with one pair, with one or two pairs in 9; intermediate
tergites withoal discal setae (rarely T3 or T4 with one small stubby adventitious setula); T5
truncate subconical without median depression, hypopygium of $ strongly prominent, surstyli
as in Text-figs 91-94.

DISTRIBUTION. Western Australia only.

Discussion. At present this genus is monotypic, containing only Chrysopasta
elegans (Macquart). It is possible, however, that this species as now understood
might be a species complex (see later discussion). Townsend (1932, 1938) considered
that elegans is generically distinct from versicolor Brauer & Bergenstamm (the type-
species of Chrysopasta) and proposed the genus Echrysopasta for elegans, but I agree
with Paramonov (1968 : 373) that elegans and versicolor are congeneric (though
specific synonymy is less certain) and therefore that Echrysopasta is a synonym of
Chrysopasta. (Paramonov cited Townsend’s name as Euchrysopasta, an incorrect
subsequent spelling.) Comparison of the types (in British Museum (Natural His-
tory)) of zabiyna Walker and elegans Macquart supports the synonymy of the
former with the latter originally indicated by Malloch (1930 : 106) and recently

REVISIONARY CLASSIFICATION OF RUTILIINI 103

repeated by Paramonov (1968 : 372), although there are some slight differences in
the $ genitalia.

The only other generic name involved in the synonymy of Chrysopasta is Roederia
Brauer & Bergenstamm, which Townsend (1938 : 412) cited as an isogenotypic
synonym of Chrysopasta with the type-species fixed by monotypy. When Brauer &
Bergenstamm (1889 : 152) first described the genus Chrysopasta with its single
species versicolor they placed the genus on its own in a family Réderiidae (‘Gruppe
XLIX’ in their classification), without at that time describing a type-genus Roederia
under this name; in their next work Brauer & Bergenstamm (1891 : 418) again
listed Chrysopasta versicolor in ‘XLIX. Gruppe Roederiidae’ but still without
a generic name Roederia. However in the next following part of their work Brauer
& Bergenstamm (1893 : 98) published the name Roederia in a key to genera and
Réderia in the generic index (op. cit. : 237), referring in the latter to the pages in
their 1889 and 1891 works on which their Gruppe Réderiidae, containing only
Chrysopasta, appeared. The name Chrysopasta does not appear in the generic key
in the 1893 work, but this genus runs down to and conforms exactly with Roederia,
_ and this fact together with the cross-references given in the 1893 index linking
Roederia to Gruppe Réderiidae (and therefore to Chrysopasta, the only contained
genus) leaves no doubt that Roederia Brauer & Bergenstamm and Chrysopasta are
one and the same genus. I therefore agree with Townsend that Roederiaisa synonym
of Chrysopasta and that versicolor is its type-species by monotypy. A possible
explanation for the confusion over the names is that Brauer & Bergenstamm, after
deciding upon the name Roederia for this Rutiliine, realized that the name was
_ preoccupied by Roederia Mik, 1881, and changed to Chrysopasta—but that the name
Roederia got published in the 1893 work by an oversight instead of Chrysopasta in
the generic key. No replacement name is, of course, required for Roederia Brauer &
Bergenstamm because of its junior objective synonymy with Chrysopasta. Para-
monov (1968) omitted Roederia completely from his treatment of Chrysopasta,
without explanation.

Paramonov (1968 : 372) implied that Chrysopasta is so different from other
_ Rutiliini when considered alongside some undescribed species known to him that it
_might be necessary later to refer the genus to some other tribe. Though these
undescribed forms to which he referred are not known to me it does not seem
_ probable on the basis of elegans that Chrysopasta could justifiably be excluded from
the Rutiliines. It possesses so many of the uniquely or at least typically Rutiliine
characteristics that it seems to me to be unquestionably a member of this tribe: for
example it shows in combination such features as more than two postalar setae,
thickly hairy suprasquamal ridge, frequently hairy prosternum and prosternal
membrane, hairy barette, dnd completely Rutiliine body facies, which together
would hardly permit it to be included in any other tribe (as least so far as the tribes
_of Prosenine Tachinidae are envisaged at present).

Within the Rutiliini the genus seems to be most nearly allied to Amphibolia, more
especially to the subgenus Paramphibolia, with which Chrysopasta possesses in general
/more common features than with the other genera. There is much similarity in
shape of the facial carina, the hairy suprasquamal ridge, the frequently strongly

104 R. W. CROSSKEY

haired prosternal regions, the head facies, and abdominal form, but Chrysopasta
differs from Amp/ibolia s.1. by having the back of the head metallic, stronger and
more extensive vestiture on the facial ridges, by the rather more slender mentum
and smaller palpi, and by the weaker abdominal chaetotaxy (in which discal setae
are absent on T3 and T4 except for the occasional occurrence of a small isolated
adventitious setula).

The head in Chrysopasta shows some unusual features not found in quite the same
form in other Rutiliini. The epistome reaches its most strongly nasute development
among the whole tribe, being extremely prominent in profile and set off from the
facial carina by a very deep concavity (Text-fig. 6), and the facial ridges are more
strongly and extensively haired above the vibrissae than in other genera; the setae
of the frontal rows also descend further down on to the extreme upper ends of the
parafacials than in other genera, so that in profile there is a smaller gap than usual
between the uppermost setulae on the facial ridges and the lowermost frontals.
The parafacial hairing is more strongly developed than is normal and extends to a
level below the bottom of the eye (the hairing virtually merging with the hairing
of the genae) ; in other Rutiliini the parafacials, when haired, have the hairing only
extending about to the bottom of the eye at most and there is a definite bare gap
between genal and parafacial hairing. The postorbits in Chrysopasta are most
characteristic, having silvery-white pollinose spots alternating with black (almost
non-pollinose) spots, this being very obvious to the naked eye and having no counter-
part elsewhere in the Rutiliines; somewhat similar pollinose areas having a shifting
chequered appearance are present on the lower ends of the parafrontals.

In many specimens the mesonotal chaetotaxy is very strong. The humeral
callus rather characteristically has only three setae set in a shallow triangle with the
innermost seta unusually strong, but there is sometimes a weakly developed fourth
seta laterad of the innermost strong one, so that the humeral chaetotaxy is basically
the same as in other Rutiliini. When several specimens are examined the chaetotaxy
is found to be much more variable than Paramonov’s (1968 : 373) description
indicates: one very strong ph (Paramonov’s ‘sublateral’) is normal, but a second may
be present ; 2 + 2 acy is common, but there may be more presutural acrostichals than
two; the post dc are usually four (length and strength very variable) ; often three or
even four supra-alars; though usually only one strong post ia seta is present the
female often has two very strong post ia. The sternopleurals are variably 1 (2) + 1
as Paramonov states. The legs of Chrysopasta are more strongly bristled than usual
in Rutiliini and the claws and pulvilli are exceedingly large, especially in the male in
which they are strikingly obvious to the naked eye; the hind tibia has a long antero-
dorsal fringe but this does not form a regular close-set comb as in many Rutiliines,
being composed instead of some fine rather hairlike setulae interposed between longer
and stronger ones so that the fringe is irregular and loose (not comb-like); the pv
apical seta of the hind tibia is much longer and stronger than usual.

The mesonotum has a bold pattern that is a little different from any other Rutiliine.
The prescutum and scutum are pale silvery bluish and have four broad bold sooty
black vittae that are discontinuous at the suture so that the scutum appears to
have a transverse row of four elongate spots of black (of which the inner pair are

REVISIONARY CLASSIFICATION OF RUTILIINI 105

shorter and more quadrate than the outer pair). The abdomen is metallic greenish
or bluish in ground colour except for the hind margins of the tergites which are
brownish black, and the paler bases of T3-T5 have an overlay of silvery or yellowish
white pollinosity which gives the abdomen a transversely banded appearance.

Paramonov (1968) drew attention to the great variability of Chrysopasta elegans,
and presuming that all the specimens so far available are truly conspecific then
elegans is the most variable species in the Rutiliini. Paramonov did not mention
the male genitalia, but these too differ greatly in different specimens (Text-figs g1—
94). This casts considerable doubt on whether elegans and its supposed synonyms
(zabirna and versicolor) are really one species, and suggests the possibility that a
species-complex is involved. In general it does not seem likely that several distinct
species occur within the relatively circumscribed area of south-west Western
Australia (where Chrysopasta is apparently confined) yet, on the other hand, it
would be very exceptional for a single species of Rutiliini (or any Tachinid) to mani-
fest the diversity in the male genitalia that is to be found amongst specimens of
Chrysopasta. The diversity in the male genitalia affects not only the size and shape
of the cerci and surstyli, but also the size of the epandrium (which in some specimens
is much bigger in relation to the cerci and surstyli than in other specimens). The
Text-figs 91-94 exemplify the diversity amongst different male specimens: Text-fig.
gi shows an enlarged epandrium, very sharply pointed surstylus and very long
tapering cerci found in a specimen from Newdegate, W.A.; Text-fig. 92 shows very
broad foliaceous surstyli and unusually small cerci found in a specimen from 6 miles
north of Watheroo, W.A.; Text-fig. 93 shows the shape of the surstyli and cerci in
the holotype of elegans (exact locality unknown); and Text-fig. 94 shows the shape
of the surstyli and cerci in the lectotype and paralectotype of zabirna from Perth,
W.A.

Differences in male genital form of this order in other Rutiliini would normally
imply distinct species and would normally be correlated with some constant, if
minor, external differences. In the case of Chrysopasta, however, there is not at
present nearly sufficient material available to determine whether the four distinct
forms of male genitalia so far detected are discrete entities or whether they represent
points along a range of variability (for example, when enough material is to hand it
might prove that there is every gradation in surstylus shape from the sharp-pointed
form of Text-fig. 91 to the broad rounded form of Text-fig. 92). The male genitalia
do not, on the basis of present knowledge, provide adequate evidence as to whether
elegans is one highly variable species or a complex of species; it therefore appears
best to retain the names zabirna and versicolor in synonymy with elegans, and to
consider all available material as conspecific, until more evidence can be adduced.

Variability exists in many external characteristics as well as in male genitalia.
_ It is possible that some of the external differences may be found to correlate with the
_ genital differences when sufficient material is studied, but no convincing correlation
has so far been discovered. Most of the variable external features show a fairly
_ complete gradation, e.g. the legs range from all reddish yellow to all brownish black
and the prosternum and prosternal membrane show all conditions from bare to very
_ strongly haired (or even bristled). Some specimens have mainly yellow pleural

106 RK. Wi. GROSSKE Y

hair and others black pleural hair (sometimes with pale hair on barette or ptero-
pleuron), and the number of median marginal setae on abdominal T3 varies in
different specimens (there is also sexual dimorphism in these setae which are often
absent in males but always present in females). The strength of the parafacial
hairing varies greatly amongst females.

It may be useful to note the condition of some of these varying features in the
primary types of elegans and zabirna in case it is later concluded that elegans is an
admixture of species. In the holotype of elegans the legs are blackish brown with
slightly more reddish tibiae, the pleural hair is black except for some golden hair on
the lower pteropleuron and the barette, and the prosternum and prosternal membrane
are strongly haired (the prosternum even bears one very long strong downwardly
directed bristle on each side) ; in the lectotype of zabirna the legs are similar to elegans
type, the pleural hair is also black but there are only a few pale hairs on the ptero-
pleuron and all the barette hair is black, and the prosternum and prosternal mem-
brane have a few strong hairs (but no definite bristle). The paralectotype specimen
of zabirna is topotypic and unquestionably conspecific with the lectotype, but has
the prosternum and prosternal membrane almost completely bare (one long recum-
bent hair on one side of the membrane, no hair at all on the prosternum itself);
otherwise it agrees with the lectotype. (Townsend, 1932 : 39, noted the two strong
prosternal setae in the holotype of elegans, as ‘2 PST’, but his statement that the
rest of the prosternum is bare is incorrect.) In the primary types of both elegans
and zabirna the prosternal hair is black. The lectotype of versicolor could not be
located for study during the present work.

The specimen from Newdegate with male genitalia as in Text-fig. gt has mainly
yellow pleural hairing (only mesopleural hair mainly black), has some long fine very
pale yellowish hair on the prosternum, and has the legs entirely reddish yellow; the
specimen from near Watheroo with male genitalia as in Text-fig. 92 has the pleural
hair similar to that of the Newdegate specimen, but has the prosternum and
prosternal membrane completely bare, and has the coxae and femora partly blackish-
brown with the rest of the legs reddish.

Macquart (1846) recorded the provenance of elegans, at the time of description,
as ‘ile Sydney’ (New South Wales), and the holotype from Bigot’s collection bears
a label by Austen recording the locality as Sydney, New South Wales; the same
type-locality was indicated in my paper on Macquart’s Australian Tachinid types
(Crosskey, 1971). It is now certain, however, beyond any real doubt that the type
must have come from Western Australia: all later known material is from Western
Australia, and it is here considered that Paramonov (1968 : 374) was right to regard
Macquart’s stated provenance as a mistake and to accept Western Australia as
being the true type-locality.

Almost nothing is known of the early stages and host-relations of Chrysopasta.
Malloch (1930 : 106) described the puparium from a specimen with its associated
adult from Swan River, and stated that another specimen from this locality had a
label reading ‘Rutilia sp. in nest of Termites, Eut. westaustraliensis’. It is extremely
improbable that Chrysopasta has a termite host as the known hosts of Rutiliini are
all coleopterous, and it should be assumed that there is no direct connection in the

REVISIONARY CLASSIFICATION OF RUTILIINI 107

life-history between Chrysopasta and termites: Townsend (1936 : 152), referring to
Malloch’s note, wrote that ‘Echrysopasta has been reared in West Australia from a
puparium found in nest of Eutermes, but it is likely that the maggot was a white
grub parasite whose host in the soil had accidentally become incorporated into the
termite nest’. This is a reasonable supposition, and the reference to a possible
inquiline relationship between Chrysopasta and termites made by Colless & McAlpine
(1970) on page 737 of The Insects of Australia must be treated with caution.

INCLUDED SPECIES

Chrysopasta elegans (Macquart). AUusTRALIA (Western Australia). [Holotype
examined].
versicolor Brauer & Bergenstamm.
zabirna (Walker). [Lectotype examined].

Genus PRODIAPHANIA Townsend

Diaphania Macquart, 1843 : 277 (120). Type-species: Diaphania testacea Macquart, 1843,
by monotypy. [Junior homonym preoccupied by Diaphania Hiibner, 1818 (Lepidoptera).]

Prodiaphania Townsend, 1927: 159. Replacement name for Diaphania Macquart, 1843,
preoccupied by Diaphania Hiibner. Type-species: Diaphania testacea Macquart, 1843, by
automatic fixation.

[Senostoma sensu authors, not Macquart. Misidentification.]

Diacnosis. Facial carina broad and flattened, sides subparallel or only slightly convergent
below, widely separating antennal bases. Epistome strongly prominent, subnasute, face in
profile deeply excavate between epistome and carina. Head of ¢ not holoptic but frons strongly
narrowed, not wider than facial carina, upper eye facets not enlarged. Parafacials haired on
upper parts. Buccal opening elongate and strongly narrowed medially, at narrowest point
subequal in width to or even a little narrower than facial carina. Genal dilation well developed.
Head pollinose, non-metallic. Arista bushy plumose or long-pubescent. Palpi exceptionally
small, sometimes papilliform, shorter than third antennal segment. Proboscis with mentum
parallel-sided or at most only a little tapering in profile. Prosternum and prosternal membrane
bare. Scutellum with apical pair of setae inserted conspicuously lower than other marginal
setae; total of four to nine pairs of scutellar marginals; disc of scutellum convex. Postalar
callus with four to six strong setae (three in holotype of cygnus). Postalar wall bare. Supra-
squamal ridge bare. Upper calypter extraordinarily developed, subequal in length in both
sexes to lower calypter. Tegula with normal long wiry posterior setulae. Costal base very
strongly explanate (wings in resting position appearing to have prominent ‘shoulders’). Ab-
dominal venter without strongly developed setae on tergites; T3 dorsally without median
marginal setae or at most with one pair in 2, marginals of T4 short; no abdominal discal setae.
T5 truncate subconical, without median depression (at most with only slight flattening or
hollowing at extreme tip).

DIsTRIBUTION. Australia only; occurring from Western Australia to Victoria and
north to Queensland, present in Tasmania.

Discussion. There has been some confusion in the nomenclature of this distinc-
tive genus, to which it is necessary to refer before considering the characteristics and
affinities. Macquart’s name Diaphania is preoccupied by Hiibner’s use of this
name in Lepidoptera, but Brauer & Bergenstamm (1889 : 126; 1893 : 175) and
Engel (1925 : 343)—being evidently unaware of the homonymy—continued to
use Diaphania Macquart as a valid name; later, however, Townsend (1927 : 159)

108 R. W. CROSSKEY

realised the homonymy and published the replacement name Prodiaphania, which
remains the valid name for this genus, and was used in publication by Malloch
(1928) : 615; 1929 : 291) and Paramonov (1968 : 355, 384). Unfortunately, the
name Senostoma Macquart has been misapplied to the genus, notably by Malloch
(1936 : 10-15) and Townsend (1938 : 426), as the result of misidentification and
erroneous synonymy of its type-species. Senostoma was described as a monotypic
genus with S. variegata Macquart as its type-species, and Macquart’s female holotype
of variegata from the Bigot collection is in the British Museum (Natural History)
(Crosskey, I97I : 291); examination of this holotype shows that vaviegata (and
therefore Senostoma) does not belong in the Rutiliini at all, but in the Prosenini
near Rhynchiodexia Bigot (of which Senostoma may well be a senior synonym).
Engel however found a female specimen in the collection of the Vienna Museum
that belonged to the species Diaphania testacea Macquart (type-species of Diaphania
= Prodiaphania) and bore a label erroneously purporting that it was Macquart’s
type of variegata, and Engel (1925 : 344) therefore synonymised variegata with
testacea; Townsend (1932 : 40) saw the same specimen in Vienna and also considered
that this synonymy was correct, recording it again later in his Manual of Myiology
(Townsend, 1938 : 426). It seems certain that neither Engel nor Townsend troubled
to compare the Vienna specimen with Macquart’s (1847 : 96) original description of
Senostoma variegata, from which it is obvious that it was not based on a Rutiliine
specimen and that Macquart’s figure (plate 5, fig. 3) scarcely resembles a Prodiaphania
species. Their mis-recognition of the type led Townsend (1938) to supplant his
name Prodiaphania with Senostoma, which would be nomenclaturally correct if the
synonymy of variegata and testacea was correct; as it is not, and as Senostoma as
correctly interpreted is not a Rutiliine, the name Prodiaphania Townsend stands
valid for the present genus.

Prodiaphania is the most distinctive and easily recognized genus in the Rutiliini,
and because of the external homogeneity of the species in conjunction with the
presence of some unusual features can be more satisfactorily defined than the other
genera. The palpi are minute (as Macquart noted in the original description of
Diaphania) and this character alone separates Prodiaphania from all other genera,
but other distinctive characters include the great enlargement of the upper calypter
(which in the wings-closed position is as long as or nearly as long as the lower calyp-
ter), the unusual elongation of the head in the epistomal axis (Text-fig. 7) with
consequent elongation of the buccal opening (Text-fig. 15), the conspicuously plumose
or unusually long-pubescent arista on which the hairing is strikingly bushy, and the
exceptionally explanate costal base (Text-fig. 26) giving the impression of basal
‘shoulders’ to the wings. Taken together these characters give Prodiaphania a very
characteristic facies and set the genus rather apart from other genera; it is not
possible to ascribe particularly close affinity to any other genus, but there is some
notable resemblance is certain features such as the elongate buccal opening and
explanate costal base to Formodexia gen. n. (but the very elongate palpi, normal
upper calypter and haired postalar wall immediately distinguish Formodexia from
Prodiaphania). The suprasquamal ridge is bare in Prodiaphania and this at once
distinguishes the genus from Chrysopasta, Amphibolia and nearly all Rutilia, but the

REVISIONARY CLASSIFICATION OF RUTILIINI 109

reduced palpi and enlarged upper calypter already referred to also provide obvious
differential characters from these genera.

The feature of enlarged upper calypter is most noteworthy, as enlargement of the
upper calypter seems to occur nowhere else in the Tachinidae. Several unrelated
Tachinids have the /ower calypter enlarged, even to such an enormous extent in the
male that the whole abdomen is covered by them, but the lower calypter is never
enlarged in Rutiliini whereas the upper one is in Prodiaphania.

The species of Prodiaphania are extraordinarily uniform in their externals, al-
though there are some minor differences in the leg chaetotaxy which can be used to
differentiate certain species or groups of species in a key. But, luckily for the
taxonomist, the male genitalia in Prodiaphania are more diverse and provide more
uniquely distinctive characters for distinguishing species than in any other genus of
Rutiliini, and both Malloch (1936) and Paramonov (1968) have made use of (and
figured) the male genitalia of several species. Some of the developments in the
genitalia of male Prodiaphania are rather bizarre (by the standards of Tachinidae)
and include the enlargement of the surstyli into enormous flattened elliptical plates
set in the transverse plane (in georgei), the development of a very large forwardly
projecting process at the base of the surstylus (as in furcata) and the development of
exceptionally dense tufts of long hair on the bases of the surstyli (as in festacea).
Sometimes the male genital characters are very distinctive even when the external
features show no evident reliable differences, and consequently dependable keys
to the species need to be based mainly on the male genitalia. Females rarely show
any reliable differences and it is difficult or well-nigh impossible to associate wild
caught females with males.

Paramonov (1968) published a key to females as well as to males, and even
described one species (walkeri) from a unique female holotype, but it seems to me
that some of Paramonov’s associations of females with males are very uncertain
and in my view it is impossible at present to identify females reliably and therefore
impossible to provide a workable key to them. Even Paramonov’s key, on assumed
associations of identity, seems very unsatisfactory, and walkeri (based only on the
female) conflicts in the key characters with those cited in the description: in walkeri
_ description (p. 400) the palpi are ‘about as long as the third antennal segment’

and the ‘calypters of about same size’, whereas the species is run out in the key

_ (p. 387) by the couplets reading ‘Upper calypter distinctly shorter than lower’ and
_ ‘Palpi half as long as third antennal segment’.
In the present work it has been concluded that Paramonov’s keys to Prodiaphania
_ are not very satisfactory, and that only a key to males is practicable at this stage.
I have therefore attempted to give here a revised key to males, based upon male
genitalia together with the few external features that seem reliable, and have given
_ figures of the male genitalia for as many species as I can (not all the types have been
to hand during this work). Before giving this key, and the list of included species,
it is necessary to allude to some of the nominal species which Paramonov (1968)
included in Prodiaphania.

Paramonov (1968 : 391) assumed that ruficornis Macquart belonged in the genus,
_ but this is incorrect; the male holotype of ruficornis has been examined and found to

IIo R. W. CROSSKEY

belong to Rutilia (Microrutilia), in which subgenus the name is a synonym of media
Macquart (see p. 90); Malloch was therefore right to place ruficornis in Rutilia and
Paramonov wrong to place it in Prodiaphamia. Regarding vittata Macquart, Brauer
(1899 : 513) was right to place it in Senostoma auct. (=Prodiaphania), and Para-
monov’s (1968 : 401) supposition that “Probably it [vittata] is a species of Rutilia
or Chetogaster’ is wrong; the female holotype (in London, not Paris as Paramonov
thought) has been examined and found to be a Prodiaphania (almost certainly the
female of testacea Macquart). Another incorrect supposition of Paramonov (1968 :
402) is that echinomides Bigot is probably not a Prodiaphania; Brauer (1899 : 512)
placed this nominal species, under the misspelling echinomyidea, in Diaphania
(=Prodiaphania) and examination of the female holotype during the present work
has shown this to be correct (though the evidence that echinomides is a synonym of
testacea as Malloch (1928c : 657) stated is not good: it appears unlikely that the
specific synonymy is correct and it is here not accepted).

The early stages and host relations of Prodiaphania are still completely unknown.

INCLUDED SPECIES

Prodiaphania arida Paramonov. AUSTRALIA (Victoria). [g paratype, data
as holotype, examined].
P. biarmata (Malloch). AusTRALia (South Australia). [Holotype examined].
P. brevitarsis Paramonoy. AUSTRALIA (New South Wales).
P. claripennis Malloch. AusTRALIA (Western Australia). [Holotype examined].
P. commoni Paramonov. AUSTRALIA (South Australia, Victoria).
P. cygnus (Malloch). AvusTRALIA (Western Australia). [Holotype examined].
P. deserta Paramonov. AUSTRALIA (New South Wales, Queensland). [J para-
type, data as holotype, examined].
P. echinomides (Bigot). AUSTRALIA (? state). [Holotype examined].
P. fullerae Paramonov. AUSTRALIA (New South Wales).
P. funebris Paramonov. AUSTRALIA (South Australia, Western Australia).
P. furcata (Malloch). Austra.ta (A.C.T., New South Wales, South Australia,
Victoria). [Holotype examined].
P. genitalis Paramonov. AUSTRALIA (A.C.T., New South Wales, Queensland,
Victoria). [Holotype examined].
paratestacea Paramonov syn. n. [g paratype, data as holotype, examined].
. georgei Malloch. AusTRALIA (New South Wales, South Australia, Western
Australia). [Holotype examined].
. minuta Paramonoy. AUSTRALIA (Queensland). [g paratype, data as holo-
type, examined].
. regina (Malloch). AUSTRALIA (Queensland). [Holotype examined].
. testacea (Macquart). AUSTRALIA (Tasmania to Queensland).
. victoriae (Malloch). AvusTRALIA (Victoria to Queensland). [Holotype
examined].
. vittata (Macquart). AUSTRALIA (South Australia). [Holotype examined:
probably 9 of testacea].
. walkeri Paramonov. AUSTRALIA (Western Australia).

Ine} as} Intec as} ns) Fa)

REVISIONARY CLASSIFICATION OF RUTILIINI III

Kery TO SPECIES OF THE GENUS PRODIAPHANIA

[Note: The following key is to males only. Females usually cannot be reliably associated

with males, and it is considered that dependable keys to females cannot be given in the present
state of knowledge; the three nominal species echinomides, vittata and walkeri cannot be included
in the key as they have female holotypes which have not yet been associated with known
males. FP. clavipennis is omitted as its male holotype has not been seen, and insufficient data
is to hand for reliable inclusion.]

I

Sternites concealed, ventral ends of tergites meeting each other in mid ventral line

of abdomen so that sternites are completely covered. Hind tibia without pd

setae. Hind femur with some pv setae basally or submedially 4 genitalia

with large foliaceous surstyli orientated in the transverse plane (Text-figs ror, 106) a
Sternites exposed partially or completely, ventral ends of tergites separated from

each other in mid venter of abdomen Hind tibia with one or more pd setae (very

rarely none). Hind femur with or without pv setae. 3 genitalia with surstyli

not of this form . : 4
Hind tibia with a secondary fringe of semi- erect setulae on the a surface in addition
to the main ad fringe. [dg genitalia not seen] . : P. commoni

[No material has been seen of this species, which is only doubtfully distinct
from biarmata]
Hind tibia without a distinct secondary a fringe in addition to the main ad fringe
(hairing of a surface all small, weak and virtually recumbent). ¢ surstyli and
cerci as Text-fig. 101 or 106 . - 3
6 genitalia with surstyli enormous in relation to Spaaidirttem and ‘sarstyli and cercl
exceedingly slender in profile (Text-fig. 101). [Western Australia]
P. georgei
6 genitalia with surstyli not exceptionally enlarged in relation to epandrium and
surstyli and cerci much broader in profile (cerci broad basally and with very

strong apical curvature) (Text-fig. 106). [South Australia] 2 : P. biarmata
Hind femur without any pv setae. Hind femur with av setae confined to “pics
third or half 2 : 5

Hind femur with some pu setae basally or - medially (very few and rather weak in
victoyiae). Hind femur with av setae and long strong hairs irregularly along its

whole length (except in victoriae, in which av setae confined to apical half) : 13
Surstylus of g genitalia with a long forwardly directed basal process that is nearly

as long as the main part of the surstylus or at least half as long (Text-figs 95, 96, 105) 6
Surstylus of 3 genitalia not of this form, if with a forwardly directed basal process

this in the form of a short bluntly rounded lobe (Text-figs 97-100, 103, 104) : 7
Basal process of surstylus rounded apically and bearing a very long dense black

hair tuft (Text-fig. 95) . . P. testacea
Basal process of surstylus slightly bifurcate and pointed apically and its very

short inconspicuous hairing (Text-figs 96 & 105). : . P. furcata
Surstylus of 3 genitalia in profile strongly curved, very long ad narrow, with sparse

hairing confined to the basal part (e.g. Text-fig. 100) : 8

Surstylus of § genitalia in profile moderately broad and only at most slightly curved,
usually (not funebris) with rather long dense neeenS on most of its length (Text-
figs 97-99, 103, 104). 9
Surstylus of ¢$ genitalia ecuenely elender me of even code along its Jength
(fig. 2C in Paramonov, 1968 : 388) : : : P. brevitarsis
Surstylus of f genitalia slightly but a. clavate, narrower near the base than on
apical half (Text-fig. 100) - P. genitalis
Surstylus of $ genitalia as in Text- fig. 103, hairing mainly eanaaca: to basal third and
consisting mostly of a moderately dense bunch of long strong hairs arising from a
swollen rounded anterobasal prominence on the surstylus. Setulae of ad fringe

112 R. W. CROSSKEY

of hind metatarsus very long and conspicuous. Hind tibia with a secondary
fringe of irregular semi-erect setulae ae the a surface in addition to the normal
ad fringe . : P. funebris
— Surstyli of 3 eee mot as in Text -fig. 103, Gates surface extensively hairy and
only the tip bare, no definite anterobasal swelling. Setulae of ad fringe of meta-
tarsus weak and inconspicuous or longer than metatarsal diameter. Hind tibia
with or without a distinct secondary a fringe of semi-erect setulae in addition to
main ad fringe . 10
to Wings without the usual eae basal daake “Ecr VSantyi of 3 genitalia as in tee
fig.97. Hind tibia without a secondary a fringe in addition to the normal ad fringe
P. minuta
— Wings with the usual sub-basal dark brown mark. Surstyli of ¢ genitalia of slightly
different form (Text-figs 98, 104, 108). Hind tibia usually with a row of distinct
small semi-erect setulae along its length which form a secondary a fringe in

addition to the usual ad fringe (setulae usually rather irregular) i II
Ir Surstyli of f genitalia rather short and broad in profile, without noticeable coreateen

(as in Text-figs 98 and 104) . 12
— Surstyli of g genitalia longer and more Seales in ae and distinctly comeeel

(Text-fig. 108) . : . P. regina

12 Hind metatarsus with ae setulae of the 75 Tow w extremely long and conspicuous,
much longer than metatarsal diameter. ¢ cerci and surstyli as in Text-fig. 104

P. deserta
-— Hind metatarsus with setulae of the ad row short and inconspicuous, not longer than
metatarsal diameter. ¢ cerci and surstyli as in Text-fig. 98 . é 5 P. arida

13 Hind femur with pv setae along the basal half. Hind femur with av setae of
irregular size along its length. Wing without sub-basal dark mark. ¢ cerci and
surstyli as in Text-fig. 107. [Western Australia] . . P. cygnus

— Hind femur with pv setae on the middle third of its length, none on the basal part.
Hind femur either with irregular av setae and strong hairs along its whole length
or with av setae confined to apical half. Wings with the usualsub-basal dark mark.
¢ cerci and surstyli as in Text-fig. 99 or Text-fig. 102. [Victoria to Queensland] 14

14 Hind femur with the pv setae very long and strong, very much longer than diameter
of femur. Hind femur with the av setae and strong hairs present all along its
length. Surstyli and cerci of $ genitalia as in Text-fig. 99, seghihe broad and
truncate and cerci not recurved at tips . ; P. fullerae

- Hind femur with the pu setae weak and few (usually =e 5 or ao sometimes
only one) and only about as long as femoral diameter. Hind femur with av setae
confined to apical half. Surstyli and cerci of ¢ genitalia as in Text-fig. 102,
surstyli blade-like with acuminate apices and cerci slightly but distinctly recurved
at tips : : > : : . : : - : : P. victoriae

Genus CHETOGASTER Macquart

Chetogastey Macquart, 1851 : 198 (225). Type-species: Chetogaster violacea Macquart, 1851,
by monotypy.

Codium Enderlein, 1936 : 417. Type-species: Rutilia oblonga Macquart sensu Enderlein (mis-
identification) [=Chetogaster violacea Macquart, 1851), by original designation.

Chaetogaster : incorrect subsequent spelling of Chetogaster Macquart, without status in nomen-
clature.

Diacnosis. Facial carina elongate subfusiform or slightly flattened above and tapering
ventrally to a sharp narrow ridge, always rather narrow. Epistome very strongly prominent,
subnasute, face in profile deeply concave between epistome and carina. Head of § not holoptic,
frons very much wider than facial carina, upper eye facets not enlarged. Parafacials bare.

REVISIONARY CLASSIFICATION OF RUTILIINI 113

Buccal opening wide in both sexes, several times broader than facial carina. Genal dilation
weakly developed, upper margin running obliquely from behind eye to lower margin of gena.
Head pollinose, non-metallic. Arista pubescent. Palpi fully developed, sexually dimorphic
(slender in males, conspicuously clavate in females). Proboscis with mentum parallel-sided in
profile. Prosternum and prosternal membrane bare. Scutellum with apical pair of setae
inserted only slightly lower than other marginal setae; total of only three pairs of marginals
(fourth supernumerary pair rarely developed); disc of scutellum convex. Postalar callus with
two strong setae. Postalar wall bare. Suprasquamal ridge bare. Upper calypter normal.

| Tegula with normal long wiry posterior setulae. Costal base not explanate (but basal costal
_ fringe sometimes strongly developed). Abdomen with marginal hair or at most only weakly

developed setae on tergite venters; T3 with one pair of erect median marginal setae ; intermediate

| tergites (T3 and T4) with one or two pairs of discal setae, these sometimes feebly developed or
_absenting. T5 truncate subconical, without median depression (at most only trace of flattening

at extreme apex).

DisTRIBUTION. Australia only, from Tasmania and South Australia to Queens-

land. Apparently unrepresented in Western Australia.

Discussion. The correct nomenclature for this genus is considered below before
giving any consideration to the characteristics and possible affinities.

Malloch (1927, 1929, 1936), Enderlein (1936) and Paramonov (1954, 1968) have
all referred to this genus by the name Chaetogaster, an incorrect subsequent spelling
of Macquart’s name Chetogaster that seems to have been first published by Brauer &
_Bergenstamm (1891 : 418, 1893 : 228); only Townsend (1932: 40, 1936 : 155,
1938 : 424) has used the correct original spelling. It is necessary to consider this
further, as both Chaetogaster and Chetogaster spellings exist in the literature for a
genus of Oligochaete Annelids. The genus Chaetogaster in Annelids was described
by Baer (1827 : 614) and the name is valid and in current use, but Gervais (1838 : 15)
cited Baer’s genus as Chetogaster and Neave’s Nomenclator Zoologicus lists Gervais’
spelling of the name with the suffixed parenthetical comment ‘(pro Chaet-Baer

1827)’—implying that Gervais’ spelling was an intentional emendation of Chaeto-
_gaster Baer. However, Gervais’ work contains no evidence that he intentionally
emended the spelling, and his use of the name Chetogaster is therefore an incorrect
subsequent spelling under Article 33 of the International Code of Zoological Nomen-
_clature; it therefore has no status in nomenclature and does not preoccupy Chetogaster
Macquart, 1851, which name stands valid for the present genus of Tachinidae.
_ Enderlein (1936) erected the genus Codium with Rutilia oblonga Macquart cited
asthe type-species. Paramonov (1968 : 365) found that Enderlein had misidentified
oblonga Macquart, and that the species that Enderlein called by this name is actually
_Chetogaster violacea Macquart. During the preparation of the present work I have
seen the specimens from Adelaide on which Enderlein based Codium, and can con-
firm that they are violacea (not oblonga), as Paramonov said; the synonymy of
Codium with Chetogaster established by Paramonov (1954) can also, therefore, be
confirmed. It should be noted that the true oblonga Macquart also belongs to
Chetogaster, and hence Codium is a synonym in any case, even if the cited nominal
species is taken to be the type.

Paramonov (1954) placed Chaetogastrina stolida Malloch, the type-species of
Chaetogastrina, in the genus Chetogaster, thereby sinking the generic name Chaeto-

114 R. W. CROSSKEY

gastrina as a synonym of Chetogaster, and he maintained this synonymy in his later
paper on the genus (Paramonov, 1968). Here it is considered that Paramonov was
in error, for stolida shows scarcely any of the diagnostic characters of Chetogaster
but—on the contrary—shows all the essential characters of Parampbolia (q.v.);
the g sternite 5 and genitalia, taken with the haired suprasquamal ridge, multiple
postalar setae, etc. show beyond doubt that the true affinities of stolida are with
assinulis Macquart, the type-species of Paramphibolia, and Chaetogastrina is here
placed as a synonym of Paramphibolia (see p. 100).

In his earlier paper, Paramonov (1954) placed three species in Chetogaster which
he later (Paramonov, 1968) removed to other genera: C. nigrithorax (Macquart) and
C. wilsoni Paramonov he placed in his genus Ola Paramonov, 1968, and C. diversa
Paramonov he placed in his genus Ruwya Paramonov, 1968. It is here agreed with
Paramonov that nigrithorax, wilsoni and diversa do not belong in the genus Cheto-
gaster; they are considered not to belong in the Rutiliini at all in the present defini-
tion of the tribe and are therefore omitted from further consideration.

The uncertainty over placement of these species is not surprising, for it is by no
means certain that Chetogaster itself should really be included in the Rutiliini.
Many of the characters of the genus fail to conform with those possessed by typical
Rutiliines and the genus should perhaps be looked upon as annectant between the
Rutiliini and the Prosenini. The narrow facial keel, robust form, haired propleuron,
and number of postalar setae (two) and scutellar marginal setae (three pairs including
the apicals) give the genus a facies reminiscent of the genus Billaea Robineau-
Desvoidy from Eurasia and Africa, and Chetogaster seems to be an Australian coun-
terpart to the African Billaea. The characters of Chetogaster certainly make it
difficult to differentiate the Rutiliini from the Prosenini in a completely satisfactory
way, but it nevertheless seems useful to maintain the two tribes in spite of the
existence of some baffiing forms that are hard to place in either tribe with confidence.
For the present I follow the traditional placement of Chetogaster and retain the genus
in the Rutiliini, pending better evidence on the real affinities (at the present time
nothing is known of the early stages and host-relations of Chetogaster).

There is no practical difficulty in distinguishing Chetogaster from other genera
within the Rutiliini as it differs from all of them by having only two setae on the
postalar callus (the normal number in Tachinidae). An interesting feature of the
genus, not found elsewhere in the Rutiliini, is the existence of sexual dimorphism
in the palpi—those of the male being slender as in other Rutiliines, but those of the
female being very conspicuously clubbed. Some Chetogaster species (three out of
the seven currently recognized species: see key) possess a small tuft of short fine
hairs on the upper part of the mediotergite just below the base of the lower calypter,
and this, too, is of unusual interest: such infrasquamal setulae (as they are usually
termed) occur in a number of unrelated Tachinidae, but are of very rare occurrence
in the whole subfamily Proseninae and occur in no other Rutiliini.

The male genitalia and sternite 5 of Chetogaster species shed no light on the affini-
ties within the Proseninae. The aedeagus is exactly similar to that of other
Rutiliini (though the membranous and the sclerotized parts of the distiphallus are
perhaps slightly longer than in most Rutiliines) and the surstyli and cerci show no

REVISIONARY CLASSIFICATION OF RUTILIINI 115

unusual features. Within the seven species three different shapes of surstylus
occur (Text-figs 45-47) and there are slight differences in the thickness of the cerci
when seen in profile: in oblonga, pellucida and argentifera (probably also auriceps of
which the male is unknown at present) the surstyli are rather long and tapering and
the cerci are shorter than the surstyli (Text-fig. 46); in canberrae the surstylus is
very small in relation to the epandrium and the cerci are much longer than the
surstyli (Text-fig. 45); and in violacea and viridis the surstyli are large, a little more
blunt than in the oblonga type, and the cerci are wide in profile (Text-fig. 47). The
three shapes are easily distinguished from one another, and it is perhaps questionable
whether the supposed species in which the male genital conformation is identical
are really distinct from one another. But on present evidence it appears best to
maintain viridis as distinct from violacea, and likewise argentifera and pellucida as
distinct from oblonga, in spite of the lack of genital differences.

Finally it may be noted here that the holotype of C. oblonga (Macquart) conflicts
with Paramonov’s (1968 : 364) placement of this species in his key: the abdomen
is not ‘reddish-yellow, semipellucid, with only a narrow, median black stripe’ as
stated in Paramonov’s key, and oblonga holotype runs to gratiosa in Paramonov’s
key. Examination of the holotypes of oblonga and gratiosa, together with other
material, indicates that the types are conspecific, and gratiosa is therefore placed
here as a new synonym of oblonga.

INCLUDED SPECIES

Chetogaster argentifera Malloch. AUSTRALIA (Australian Capital Territory,
New South Wales, Victoria). [Holotype examined].

C. auriceps Paramonov. AUSTRALIA (Queensland).

C. canberrae Paramonov. AUSTRALIA (Victoria to Queensland). [Holotype
examined].

C. oblonga (Macquart). AUSTRALIA (New South Wales, Victoria). [Holotype
examined].

gratiosa Paramonov syn. n. [Holotype examined].
C. pellucida Paramonov. AvsTRALIA (New South Wales). [Holotype exa-

mined].

C. violacea Macquart. AUSTRALIA (Tasmania to Queensland). [Holotype
examined].

C. viridis Malloch. AustRaLia (New South Wales, Queensland). [Holotype
examined].

Key To SPECIES OF THE GENUS CHETOGASTER

1 Calyptrae bright orange-yellow. Wings conspicuously yellow or yellow-orange
anterobasally (as all basal veins with this colour) and faintly but distinctly tinged
with yellowish or smoky brownish colour elsewhere. Infrasquamal setulae
present.? g surstyli and cerci in profile as in Text-fig. 47 (cercus rather broad and
sharply narrowing at tip). Large species, length normally 16-20 mm. , : 2

3 The infrasquamal setulae consist of a small tuft of hairs immediately below the base of the lower
calypter on the mediotergite. They are very small and careful examination is required.

116 R. W. CROSSKEY

Calyptrae not yellow, either white or partly to completely smoky brown. Wings
entirely clear hyaline with dark veins. Infrasquamal setulae absent (except in
canberrae). § surstyli and cerci not of this shape, cerci more slender (Text-figs 45 &

46). Smaller species, length normally io-16mm _ . 3

Abdomen metallic green to blue-violet on dorsum and siuleeeons on cea spdouseel
venter with scarcely any trace of whitish pollinosity visible to naked eye in any
light. Mesonotum dark green to blue-green with the pollinosity distinctly tinged
with yellow. Scutellum green to violet. g vertex o-12-0-14 of head-width.
Abdominal T5 with unusually strong, often distinctly spiniform, bristling C. viridis

Abdomen bluish black with purplish tinge dorsally and deep violet or reddish violet
ventrally; venter with white pollinosity extremely conspicuous to naked eye when
abdomen viewed from behind. Mesonotum very dark blue-black or violaceous
black with the pollinosity white. Scutellum dark violet (sometimes with brownish
tinge) to purplish black. ¢ vertex consistently wider, 0:15-0:17 of head-width.
Abdominal T5 with strong bristling but the setae long and not at all spiniform

C. violacea

Palpi and epistome black or brownish black. Upper and lower calypter both dark
smoky brown. Infrasquamal setulae present (but minute and inconspicuous).

6 genitalia with surstylus shaped as in Text-fig. 45, cerci much longer than the
relatively small surstyli . : - C. canberrae

Palpi and epistome yellow or tawny eellewe Crore is or Bees white, if
largely smoky brown then at least basal half of upper calypter white. Infra-
squamal setulae absent. ¢ genitalia with ae pee SS a as in Text-fig. 46, cerci
shorter than surstyi . 4

Ground colour of abdomen aan yallowih or very pale cedcice bean eh a
conspicuous black median vitta (abdomen distinctly pallid to naked eye on either
side of the black mid-line). Scutellum dark brown with little or no trace of
metallic colour. : C. pellucida

Ground colour of abdomen very dace metallic preeaisn or ooo appearing dark
brownish in some lights only, darkest mid dorsally but without a definite black vitta.
Scutellum distinctly metallic steely blue-black or purplish violet 3

Third antennal segment uniformly bright orange. Parafacials bright aaa
pollinose. Ground colour of parafrontals tawny reddish. Calyptrae almost all
white, only the border darkened. Abdomen with pollinosity of the dorsum almost
entirely whitish (at most only yellowish brown near the mid-line)

C. argentifera, C. auriceps
[C. auriceps is known only from the 2 holotype which has not been examined.
Paramonov (1968) distinguished it from argentifera mainly by the yellow pollinose,
instead of silvery whitish pollinose, parafrontals. It appears doubtful whether
auriceps is specifically distinct from argentifera.]

Third antennal segment partly or mainly brown, at least infuscate along one edge.
Parafacials mainly whitish pollinose (if lower parts somewhat yellowish pollinose
as in some @ specimens then calyptrae largely smoky brown). Ground colour of
parafrontals blackish. Calyptrae at least faintly infuscate over much of their
surface or lower calypter broadly darkened against hind edge, calyptrae of 9?
usually dark smoky brown except for basal half of upper calypter. Abdomen with
pollinosity of the dorsum mainly dark yellowish brown or coppery brown on the
median third (the darker pollinosity of the middle of the abdomen contrasting with
the whitish lateral pollinosity in some lights) . : . . : C. oblonga

LECTOTYPE DESIGNATIONS

Lectotypes are newly designated in this section for the 29 species-group taxa of

Rutiliini which were originally based upon syntypes and for which no previous valid

REVISIONARY CLASSIFICATION OF RUTILIINI 1a)

lectotype designations or restrictions have been published; the lectotypes have been
appropriately labelled. The names are listed in alphabetical order under their
original generic assignments.

For some taxa not all of the original syntypes have been located and examined,
but all available syntypes other than the lectotypes (i.e. the paralectotypes) are
noted in the following list and have been labelled as paralectotypes.

It should be noted that several of Enderlein’s types have labels in his hand-
writing on which the generic name is different from that with which the specific
name was combined in publication, and attention has been drawn to such discrepan-
cies where they occur by quoting the labels. Enderlein (1936) evidently discovered,
while his paper was in press, that his genus Habrota was based upon the same type-
species as Chrysorutilia Townsend, 1915, and that some of his new species should be
assigned to Townsend’s genus; thus these were published in Chrysorutilia but
remained labelled as Habrota; but, apart from this, there are instances where Ender-
lein seems simply to have changed his mind on the generic position of some new
species between the time of labelling the types and publishing the paper. However,
despite these discrepancies between label name and published name, there is never
any doubt about the identity of the types for any nominal species-group taxon.

Chrysorutilia angustigena Enderlein, 1936 : 403.

Described from two g syntypes from the same type-locality. LECTOTYPE. AvustTRALia,
Queensland, Herberton, i.1910 (J. P. Dodd) (in Museum fiir Naturkunde der Humboldt-Univer-
sitat, Berlin). The lectotype bears Enderlein’s name label ‘Habrota angustigena Type Ender.

ie

Chrysorutilia caeruleata Enderlein, 1936 : 402.

Described from ¢ and 2 syntypes (total number not clearly stated) from the same type-locality
on different dates. LECTOTYE g. Avustrraria, Western Australia, Marloo Station, ix.1935
(A. Goerling) (in Museum fiir Naturkunde der Humboldt-Universitat, Berlin). Paralectotypes:
23, 19, same data as lectotype except dates ix. 1934 for one g and ii.1935 for 2) ($2 in MNHU,
Berlin; § with date ix.1934 in BMNH, London).

__ The lectotype bears Enderlein’s name label ‘Habrota caeruleata Type Enderl.. g’, and the
_ paralectotypes cited also bear Enderlein labels with the ‘Habrota’ generic name. The 2
paralectotype is probably mis-associated with the lectotype.

Chrysorutilia caesia Enderlein, 1936 : 402.

__ Described from one g and one 2 syntypes from ‘Australien’ and four 9 syntypes from Queens-
| land, of which the ¢ is designated. LECTOTYPE g. Avustravia (Clement) (in Museum fiir
_ Naturkunde der Humboldt-Universitat, Berlin). Paralectotype: one 2, same data as lectotype

(MNHU, Berlin).

| Enderlein’s name label is lost from the lectotype specimen (though its identity as the original
| 6 syntype is certain from the label ‘Australien Clement’ and agreement with description), but
the paralectotype 2 has Enderlein’s name label ‘Rutilia caesia Type Enderl. 9’.

Chrysorutilia goerlingiana Enderlein, 1936 : 404.

Described from two ? syntypes from the same type-locality. LECTOTYPE 92. AusrTrRatia,
| Western Australia, Wurarga, Marloo Station, i.1935 (A. Goerling) (in Museum fiir Naturkunde
der Humboldt-Universitat, Berlin). The lectotype bears Enderlein’s name label ‘Rutilia
_ goerlingiana Type Enderl. 9’.

118 R. W. CROSSKEY

Chrysorutilia lineata Enderlein, 1936 : 407.

Described from one g and one 2 syntypes from the same locality. LECTOTYPE 3. AustrRa-
L1A, Western Australia, Wurarga, Marloo Station (A. Goerling) (in Museum fiir Naturkunde der
Humboldt-Universitat, Berlin). The lectotype bears Enderlein’s name label ‘Habrota lineata
Type Enderl. Q’.

Chrysorutilia rufibarbis Enderlein, 1936 : 405.

Described from one 2? syntype from ‘Nord-Australien’, and one g and one 2 syntypes from
Western Australia, of which the g is here designated. LECTOTYPE. Austratia, Western
Australia, Wurarga, Marloo Station, ix—x.1934 (A. Goerling) (in Museum fiir Naturkunde der
Humboldt-Universitat, Berlin). Paralectotype: one 9, same data as lectotype except date
x—xi.1934 (MNHU, Berlin).

The lectotype bears Enderlein’s name label ‘Habrota rufibarbis Type Enderl. g’ and the year
date is 1934 (not 1935 as Enderlein cited). The paralectotype 9 (which is much larger than,
and perhaps not conspecific with, the lectotype) also has ‘Habrota’ on Enderlein’s name label.

Chrysorutilia splendida var. confluens Enderlein, 1936 : 407.

Described from one 2 syntype from Western Australia, one 2 syntype from Sydney, and ag
and five 2 syntypes from Queensland. LECTOTYPE. AusTRALia, Queensland (in Museum
fiir Naturkunde der Humboldt-Universitat, Berlin). Paralectotype: one 9, AUSTRALIA, Western
Australia, Swan River (Ddmel) (MNHU, Berlin).

The lectotype bears Enderlein’s name label ‘Habrota splendida (Donov. 1805) var. confluens
Type Enderl. 3’. The 9 paralectotype (which has pale pleural hair and is probably wrongly
associated) is labelled ‘Rutilia splendida Guér. var. confluens Type Enderl. 9’.

Dexia zabirna Walker, 1849 : 863.

Described from two $ syntypes with identical data from the British Museum collection, each
of which bears an old accession label reading ‘47 109’ on one side and ‘W. Austr-alia’ on the other,
and a pencilled locality label in Austen’s handwriting. LECTOTYPE. AustTRa.ia, Western
Australia, Perth (G. Clifton) (in British Museum (Natural History), London). Paralectotype:
one g, same data as lectotype (BMNH, London).

Donovanius fulgidus var. nigribasis Enderlein, 1936 : 411.

Described from an unstated number of g and 2 syntypes from the same locality. LECTO-
TYPE 3. AuvsTRa.ia, Western Australia, Wurarga, Marloo Station, 6.vi.1935 (A. Goerling) (in
Museum fiir Naturkunde der Humboldt-Universitat, Berlin). Paralectotype: one 9, same
data as lectotype except date ix.1935 (MNHU, Berlin).

Idania ralumensis Enderlein, 1936 : 409.

Described from two 9 syntypes with the same data. LECTOTYPE®?. Papua NEw Guinea,
New Britain, Ralum, 30.viii.1896 (Ff. Dahl) (in Museum fiir Naturkunde der Humboldt-Univer-
sitat, Berlin). The lectotype bears Enderlein’s name label ‘Rutilia ralumensis Type Enderl. 9’.

Laccura engeli Enderlein, 1936 : 431.

Described from two 9 syntypes with the same data. LECTOTYPE 92. INnponeEsiA, Banda
Islands, Run [=Roon] Island (south of Ceram) (in Museum fiir Naturkunde der Humboldt-
Universitat, Berlin).

Pancala eos Enderlein, 1936 : 423.

Described from three ¢ and seven 9 syntypes from Celebes. LECTOTYPE <3. INDoNgEsiA,
Celebes, Bonthain, Wawa Karaeng, 1100 m, viii.1931 (G. Heinrich) (in Museum fiir Naturkunde
der Humboldt-Universitat, Berlin). Paralectotypes: one g, two 9, same data as lectotype
($ in BMNH, 92 in MNHU); one 9, same data as lectotype except date ix—x.1931 (MNHU).

Pancala gemmata Enderlein, 1936 : 424.
Described from one 2 and two g syntypes from New Britain. LECTOTYPE g. Papua
New Guinea, New Britain, Ralum, Karakaul, 30.viii.1896 (F. Dahl) (in Museum fiir Natur-

REVISIONARY CLASSIFICATION OF RUTILIINI 119

kunde der Humboldt-Universitat, Berlin). The lectotype bears Enderlein’s name label
‘Chromocharis gemmata Type Enderl. 3’.

Pancala gemmata var. viridescens Enderlein, 1936 : 424.

Described from two g syntypes with the same data. LECTOTYPE. Papua NEw GUINEA,
New Britain, Kinigunang (C. Ribbe) (in Museum fiir Naturkunde der Humboldt-Universitat,
Berlin). The lectotype bears Enderlein’s name label ‘Pancala gemmata n. sp. var viridescens
Type Enderl. 3’.

Prosenostoma nigripes Enderlein, 1936 : 435.

Described from two 3 syntypes from Queensland, one from Cairns and the other from Herber-
ton. LECTOTYPE 3. AvustrRAtia, Queensland, Herberton, 3700 ft, ii.1g911 (Dodd) (in Museum
fiir Naturkunde der Humboldt-Universitat, Berlin). The lectotype bears Enderlein’s name
label ‘Agalmia nigripes Type Enderl. 3”.

Psaronia bisetosa Enderlein, 1936 : 414.

Described from two 9 syntypes from Western Australia. LECTOTYPE 9. AustTRALia,
Western Australia, Wurarga, Marloo Station, 1935 (A. Goerling) (in Museum fiir Naturkunde
der Humboldt-Universitat, Berlin).

Pseudoformosia pauper de Meijere, 1904 : 178.

Described from three 3 syntypes collected by Bernstein in northern Halmahera. LECTO-
TYPE 9. Invonesia, Moluccas, N. Halmahera (Bernstein) (in Rijksmuseum van Natuurlijke
Historie, Leiden). Paralectotype: one gj, same data as lectotype (in British Museum (Natural
History), London).

The syntypes cited above each bear a printed label reading ‘Bernstein Noord Halmaheira’.
The Leiden Museum collection contains, standing with the lectotype, two § specimens each with
a similar printed label reading ‘Bernstein Morotai’ but these specimens have no type-status.
Similarly, the collection of the Zodlogisch Museum, Amsterdam, contains one ¢ and one 9
specimen of pawper with the printed ‘Bernstein Morotai’ labels, but these specimens also have
no type-status.

Rutilia atribasis Walker, 1861b : 288.

The type-material consists of two conspecific 2 syntypes, though the number of specimens
was not stated by Walker. LECTOTYPE 9. Inponesta, Moluccas, Batjan [publ. as ‘Batch-
ian’] (A. R. Wallace) (in British Museum (Natural History), London). Paralectotype: one 9,
same data as lectotype (in University Museum, Oxford).

Rutilia desvoidyi Guérin-Méneville, 1843 : 269.

Described from three syntypes (two 3, one 9) from ‘Nouvelle-Hollande’, of which the males
have not been located and are presumed lost and of which the 2 syntype (here designated) is in
Paris. LECTOTYPE. Auvsrratia (no other data) (in Muséum National d’Histoire Naturelle,
Paris, standing in Macquart collection under reference number 2310).

The lectotype bears a label in Guérin-Méneville’s writing which reads ‘Rutilia Desvoidyi.
guer. R. vivipara. Rob. D. p. 321.3. non vivipara f. N.elle hollande’, this label making reference
to the fact that Guérin-Méneville considered that Rutilia vivipava of Robineau-Desvoidy was a
misidentification of the true R. vivipava (Fabricius). There is no known evidence to support
Guérin-Méneville’s supposition, and as the lectotype of desvoidyi is conspecific with the neotype
of Rutilia vivipara (Fabricius) (herein designated) the name desvoidyi falls as a synonym of
vivipara.

Rutilia inornata Guérin-Méneville, 1843 : 268.

Described from five 2 syntypes from “Nouvelle-Hollande’ recorded in the original description
as ‘5 individus femelles provenant du méme marchand, 2 dans la coll. du Muséum, 3 dans la
mienne’; the museum referred to is the Natural History Museum in Paris. The two syntypes
in Paris Museum still exist there as part of the Macquart collection, but the other three syntypes

120 R, W. CROSSKEY

have not been located and are presumed lost. LECTOTYPE 9. Austratia (no other data)
(in Muséum National d’Histoire Naturelle, Paris, standing in Macquart collection under refer-
ence number 2311). Paralectotype: one 2, same data as lectotype (MNHN).

Both lectotype and paralectotype are in bad condition, the latter consisting only of the thoracic
shell and right wing; the lectotype is eaten out in places, lacks antennae and mouthparts and
has been glued in places. The lectotype bears a label in Guérin-Méneville’s writing which reads
‘Rutilia inornata. guer. ic. R. a. (type) Nelle holl.’.

Rutilia lucigena Walker, 1861a : 242.

Described from both sexes but number of specimens not stated, though actually consisting
of three conspecific syntypes. LECTOTYPE 4. INponEs1AN NEw Guinea, Dorey
(A. R. Wallace) (in British Museum (Natural History), London). Paralectotypes: onedg, one 9,
same data as lectotype (BMNH).

Rutilia pretiosa Snellen van Vollenhoven, 1863 : 15.

Described from one g and one 2 syntype, both still in Leiden Museum. LECTOTYPE 4.
InpDoneEs14, Moluccas, Ternate (Bernstein) (in Rijksmuseum van Natuurlijke Historie, Leiden).
Paralectotype: one g, same data as lectotype except collector Forsten (also in RMNH, Leiden).

The lectotype bears a circular white label with the words ‘Bernst. Ternate’ in ink, and the
paralectotype bears a similar label with the words ‘Forsten Ternate’.

Rutilia pubicollis Thomson, 1869 : 530.

Described from an unstated number of § specimens from ‘Sidney’; the type-material actually
consists of two identically labelled and set syntypes in Stockholm. LECTOTYPE. AustRa-
L1A, New South Wales, Sydney (Kinb.) (in Naturhistoriska Riksmuseum, Stockholm). Para-
lectotype: one 3, same data as lectotype (NR, Stockholm).

Each of the two syntypes has a small rectangular printed label reading ‘Sidney’ and another
rectangular printed label reading ‘Kinb.’.

Rutilia saturatissima Walker, 1861b : 287.

Described from both sexes without statement of the number of specimens. Type-material
consists of one syntype of each sex (conspecific) LECTOTYPE $. Inponesia, Moluccas,
Batjan [publ. as ‘Batchian’] (A. R. Wallace) (in British Museum (Natural History), London).
Paralectotype: one 9, same data as lectotype (BMNH).

Rutilia speciosa Erichson, 1842 : 273.

Erichson did not state the sex of the original material or the number of specimens, but
Enderlein (1936 : 430) showed that there were three 9 syntypes collected by Schayer in Tas-
mania which he referred to as ‘die Typen Erichson’s’. Two of these syntypes have been seen.
LECTOTYPE 2. AusrTRALta, Tasmania (Schayer) (in Museum fiir Naturkunde der Humboldt-
Universitat, Berlin). Paralectotype: one 2, same data as lectotype (MNHU, Berlin).

The lectotype bears two rectangular blue labels with the handwritten words ‘Vandiem.
Schayer’ and ‘Speciosa Er.’ respectively (these probably both in Erichson’s hand), and also a
yellow rectangular label with the printed number ‘3554’. The paralectotype bears a label
reading “Tasmanien Schayer’ in Enderlein’s writing. The third syntype has not been seen
but is probably still in the MNHU, Berlin, collection.

Rutilia spinipectus Thomson, 1869 : 530.

Described from an unstated number of specimens of both sexes from ‘Sidney’; the type-
material consists of one ¢ and one 2 syntype. LECTOTYPE g. Austraria, New South
Wales, Sydney (Kinb.) (in Naturhistoriska Riksmuseum, Stockholm). Paralectotype: one 9,
same data as lectotype (NR, Stockholm).

Each of the syntypes has a small rectangular printed label reading ‘Sidney’ and another
rectangular printed label reading ‘Kinb.’.

Rutilia volucelloides Walker, 1861b : 289.
Walker described this species from both sexes, but only a single syntype now exists; this is

REVISIONARY CLASSIFICATION OF RUTILIINI 121

here designated as lectotype. LECTOTYPE g. Inponesia, Moluccas, Batjan [publ. as
‘Batchian’] (A. R. Wallace) (in British Museum (Natural History), London).

Senostoma flavipes Brauer & Bergenstamm, 1889 : 126 (58).

Brauer & Bergenstamm described both the g and the @ of this species, and the original material
consisted therefore of at least two syntypes; the locality was cited as ‘Neuholland’. Engel
(1925 : 375) saw the original material and referred to it as ‘g and 2 Neu Holland (Typen No. 38
u. 39)’, clearly indicating that the syntypes consisted of one of each sex. The syntype cannot
now be found in the Vienna Museum collection, but the 9 syntype is still present there: it is here
designated as lectotype. LECTOTYPE 9. [‘New HoLtranp’] (in Naturhistorisches Museum,
Vienna).

The lectotype has a square white label inscribed in slightly faded ink ‘Dup. n 39’, the number
39 conforming with the number cited for the 2 syntype by Engel.

Townsend (1932, 1938) misunderstood the type-material of flavipes and the specimens from
Western Australia cited by him as ‘Holotype’, and in his notes in the U.S. National Museum,
Washington, as ‘Holotype’ and ‘Paratype’ have no type-status (see further discussion of this
in the treatment of the subgenus Microrutilia).

Direct comparison of the lectotype ? of flavipes Brauer & Bergenstamm with the lectotype 2
of fulviventris Bigot shows that the types are in perfect agreement and unquestionably conspeci-
fic. Hence the name flavipes is sunk as a new synonym of fulviventris (see p. 90).

Tachina inusta Wiedemann, 1830 : 306.

Described from an unstated number of specimens of unstated sex from ‘Port Jackson in
Neuholland’ (i.e. Sydney), and recorded by Wiedemann as in the Berlin Museum. Enderlein
(1936 : 412) found four ? specimens collected by Eschscholtz at Port Jackson which he recorded
as ‘Typen Widemann’s’, of which two have'been examined and one here designated as lectotype.
LECTOTYPE 2. AvustTrRaLia, New South Wales, Sydney [publ. as ‘Port Jackson’] (Eschscholtz)
(in Museum fiir Naturkunde der Humboldt-Universitat, Berlin). Paralectotype: one 9, same
data as lectotype (MNHU, Berlin).

The lectotype bears a rectangular faded bluish-grey label with the words ‘P. Jackson. Esch.’,
and another faded bluish grey label reading ‘inusta Tachin. inusta Wied.’, the writing on both
labels probably being Wiedemann’s (as it closely resembles the writing on Wiedemann’s type
labels in Copenhagen); in addition the lectotype has a printed label ‘3553’. The paralectotype
is labelled ‘Australien Port Jackson Eschscholtz’ in Enderlein’s writing.

NEOTYPE DESIGNATIONS

The types of some of the Rutiliini described by the early workers (i.e. before 1850)
are lost and have never been reported in the literature as having been seen by
anyone since the time of description. The type-material is considered to be cer-
tainly lost in the case of fifteen nominal species described by Donovan, Fabricius,
Guérin-Méneville, Macquart, Robineau-Desvoidy and Walker. Most of the names
involved apply to, or are believed on all available evidence to apply to, distinct
species and to be taxonomically valid; several of them have been in use for many
years, but have not always been applied to the same species. Some of the supposedly
single species to which some of these old names have been applied are now known
to be complexes of very closely allied species, and to disentangle past confusion (as
well as to obviate the possibility of future misidentifications) it is considered highly
desirable to establish neotypes so that the meanings of the names can be fixed.

Opinions vary among taxonomists on the desirability of neotypes: some workers
maintain that they are scarcely, if ever, justified and others regard them as desirable
for every case where there is no surviving original type-material. I take an inter-

122 R. W. CROSSKEY

mediate viewpoint and regard them as desirable in groups that have been much
confused with diverse applications of the same names (with a high potential for
continuing future confusion), but only for names that are maintained as valid or are
unfamiliar synonyms for which there is good reason to have neotypes. In the present
work, neotypes are established for twelve of the fifteen names for which original
type-material no longer exists; the other three names (australasia Gray, fulvipes
Guérin-Méneville, and vidwa Guérin-Méneville) have been accepted for many years
as junior synonyms, there are no reasons for doubting the rightness of the synonymies,
and neotypes are not therefore established for these names.

Special care has been taken here to ensure that the neotypes proposed are validly
designated according to the rather stringent terms of Article 75 of the International
Code of Zoological Nomenclature (1961). The present paper contains the results of
revisory work based upon a study of the whole tribe Rutiliini, during which it has
been found necessary for neotypes to be established in the interests of stability of
nomenclature, by resolving confused and doubtful identities (Article 75 (a) (i).
For each neotype it is considered that the qualifying conditions specified in Article
75 (c) (1-6) are fully satisfied in the present work for the following reasons (in order
of the six qualifying conditions): (1) The placement of the species for which the
neotype stands in a key to all the species of its higher taxon, together with a figure
of the male genitalia and a bibliographic reference to the original description,
constitute a statement of the characters which (in my view) differentiate the species
for which the neotype is designated; (2) The data is published here for each neotype
and each has been appropriately labelled; (3) Original material has been personally
searched for in all likely museum collections, none has been found and none has been
mentioned by any author since the time of description, from which facts it is con-
cluded that all original type-material of each nominal species for which a neotype is
designated is either lost or destroyed; (4) The characters of each neotype are com-
pletely consistent with the original description, except that a male specimen has
usually been chosen as neotype (for the male has better characters than the female)
even though the original description or figure was based or apparently based on a
female specimen; (5) The exact provenance of the original material is usually uncer-
tain, in which case the neotype specimen is chosen from an area which is considered
a likely or possible original provenance having regard to all the circumstances of the
time when the species was described (e.g. early coastal development of Australia) ;
(6) The museum depository is named for each neotype (the neotypes are deposited
either in the Australian National Insect Collection at Canberra or in the British
Museum (Natural History) in London, which maintain research collections and
make types available for study).

The twelve neotypes designated are all for nominal species belonging in the genus
Rutilia Robineau-Desvoidy s.1. ; ten of them (viz. chersipho, decora, formosa, imperialis,
lepida, pellucens, regalis, setosa, splendida and vivipara) apply to nominal species of
which the names are considered taxonomically valid in the present work, and the
other two (durvillei and serena) apply to newly established synonyms. The twelve
nominal species belong in three subgenera of Rutilia and their neotypes therefore
show the subgeneric characteristics which define the subgenera (see subgeneric

REVISIONARY CLASSIFICATION OF RUTILIINI 123

diagnoses): chersipho, decora, formosa, imperialis and splendida belong in the sub-
genus Chrysorutilia Townsend; lepida, pellucens and regalis in the subgenus Dono-
vanius ; and setosa and vivipara in Rutilia s.str.; serena is considered to be a synonym
of rubriceps, a species of Chrysorutilia, and durvillei is a synonym of vivipara (the
type-species of Rutilia).

The names of the nominal species for which neotypes are designated in the follow-
ing list are arranged in alphabetical order of their original binomina; data and
depository of each neotype are cited first, followed by any appropriate annotation.

Dexia chersipho Walker, 1849 : 864.

NEOTYPE ¢g. AvusTRALiA, Western Australia, Waroona, 21.xi.1908 (G. F. Berthoud) (in
British Museum (Natural History), London).

This species was described from a specimen of unknown locality in Children’s collection.
The specimen is not among those in the BMNH from Children’s collection, has never been
located and appears certainly lost. Walker’s description of the very distinctive colour pattern
and his placement in the Rutilia group of Dexia fit perfectly the common Western Australian
species later called evichsonii Engel, 1925, and there is considered to be no doubt about the iden-
tity of chersipho: it is an older name for evichsonii, which is herein placed in new synonymy with
chersipho.

Dexia serena Walker, 1849 : 865.

NEOTYPE®. Inp1a, Maharashtra, Purandhar, near Poona, c. 3000 ft, on bush (E. P. Sewell)
(in British Museum (Natural History), London).

Described from a female specimen from Madras presented to BMNH by Walter Elliott but
now lost. The description clearly indicates that this is the only species of Chrysorutilia so far
described from India, and fits exactly with the holotype female of Rutilia nitens Macquart,
1851, from India. Enderlein (1936 : 403) placed nitens as a synonym of serena and this synony-
my is here upheld, the primary types being undoubtedly conspecific. In the present revisionary
work, however, no differences have been found between the Indian material and Rutilia rubriceps
Macquart from Australia, so that both serena and nitens are herein treated as new synonyms of
vubriceps.

Musca splendida Donovan, 1805 : plate figure, unpaginated description.

NEOTYPE 3. Avustratia, Queensland, Brisbane, 8.xi.1907 (A. J. C. Wightman) (in British
Museum Natural History), London).

Townsend’s (1932 : 39; 1938 : 416) statement of ‘Ht [=holotype] in London’ for splendida
is based on the mere supposition that Donovan’s type must be in the British Museum (Natural
History). In fact there is no such specimen in BMNH collection, and no original material has
ever been located. Donovan’s figure was based on a specimen from McLeay’s cabinet from New
South Wales, but the specimen has not been found in the McLeay Museum, Sydney. Donovan’s
coloured plate of splendida shows, at natural size, a large metallic blue-green Rutilia species
with yellow head and metallic spots on the abdomen, clearly a species of Chrysorutilia. The
name splendida as used in the literature refers to a complex of three species characterized by
three quite distinct forms of male genitalia, one of which was figured by Malloch (1929 : 296)
under the name splendida. As the species figured by Malloch is apparently the commonest
of the three, occurring widely in New South Wales and Queensland (and therefore the most
likely to have been seen by Donovan), it is from this species that the neotype has been desig-
nated and to which the name sPlendida is therefore restricted.

Malloch (1928c : 660) and Townsend (1938 : 416) cited the publication date of splendida as
1798 in error. The correct date is 1805, as cited by Enderlein (1936 : 407).

124 R. W. CROSSKEY

Rutilia decora Guérin-Méneville, 1843 : 266.

NEOTYPE g. AvusTRALIA, Queensland, Burpengary (T. L. Bancroft) (in British Museum
(Natural History), London).

This species was described from a single female from ‘Nouvelle-Hollande’. The holotype is
lost, and Townsend’s (1932 : 38) statement of a female holotype in Paris is in error: there are
no specimens remaining in Paris of decora among Guérin-Méneville’s material (which is scattered
in Macquart’s collection), but there are specimens standing in Macquart’s collection under the
name decova which are later determined material. The description of decora is very full and
detailed and certainly applies to a species of the splendida-complex of Chrysorutilia, and Malloch
(1928c : 660), Townsend (1932 : 38-39) and Enderlein (1936 : 407) treated decora as a synonym
of splendida (though Enderlein accorded it varietal status). In the present revision it is con-
sidered best to apply the name to a species which is very closely allied to splendida but has
distinctly different male genitalia (see key and Text-figures); this recovers the name decorva
from synonymy and obviates the need to add a new name to the literature. It is considered
far better to bring the old name decora into use for a valid species (which exactly fits the original
description) than to describe a new species unnecessarily.

Rutilia durvillei Robineau-Desvoidy, 1830 : 321.

NEOTYPE g. Austraria, New South Wales, Hawkesbury River, 28.xi.1914 (in British
Museum (Natural History), London).

Robineau-Desvoidy described this nominal species from a male specimen collected at Port
Jackson (i.e. Sydney harbour) by Captain Dumont-Durville and given to him by Guérin-
Méneville. The neotype specimen is from the Hawkesbury River, which enters the sea just
north of Port Jackson. Robineau-Desvoidy’s description refers to a specimen closely resembling
vivipava Fabricius and having a bluish trace on the thoracic dorsum and a series of stiff bristles
on each abdominal segment (‘une série de poils raides et noirs au sommet de chaque segment’).
This description makes it clear that the type had conspicuous bristling on the third abdominal
tergite (apparent second segment) as well as on the other segments, and from this statement
(plus the remainder of the description) it seems certain that the name durvillei alluded to a
species of the subgenus Rutilia s. str. In this subgenus only vivipara (the type-species) has the
abdominal bristling as bold and conspicuous as Robineau-Desvoidy’s description implies, and I
believe it to be almost certain that the durvillei type was a specimen of vivipara (some speci-
mens of the latter have a slight bluish or purplish tinge on the scutum). I have therefore
designated as neotype of durviilei a male specimen of vivipara (a specimen considered unquestion-
ably conspecific with the neotype of vivipava herein designated also); the name durvillei is
therefore disposed of in synonymy with vivipara, which is a better course than bringing the
name into use on shaky evidence for some valid species.

Rutilia formosa Robineau-Desvoidy, 1830 : 320 (name attributed to ‘Donavan’ in error).

NEOTYPES AustrRa.ia, New South Wales, near Lake George, 25.xi.1953 (S. J. Paramonov)
(in Australian National Insect Collection, Canberra).

Robineau-Desvoidy described this species from a specimen of unstated sex that stood in
Count Dejean’s collection from ‘Nouvelle-Hollande’. Under his name Rutilia formosa,
Robineau-Desvoidy attributed the name to Donovan with the statement ‘Musca formosa,
Donavan’ and after the description wrote ‘Donavan l’a décrite et figurée parmi les insectes
remarquables de ce pays’ (i.e. Australia). It appears that Robineau-Desvoidy wrote ‘foymosa’
in error, as Donovan described and figured only one species of Rutiliine, namely Musca splendida;
there is no such nominal species as foymosa Donovan. Although Robineau-Desvoidy’s descrip-
tion may have been intended as a redescription of splendida Donovan there is no conclusive
evidence of this, and (in common with other workers on the group) I therefore accept Rutilia
formosa Robineau-Desvoidy as a nominal species originally based upon a type-specimen. This
type-specimen, along with all the other Diptera from Count Dejean’s collection, is now lost and
there are no specimens in Robineau-Desvoidy’s collection or any other authoritatively identified
specimens by which the identity of foymosa R.-D. can be determined. Robineau-Desvoidy’s

REVISIONARY CLASSIFICATION OF RUTILIINI 125

description, however, is clearly that of a species of the subgeneric concept Chrysorutilia and
contains certain clues on which species he must have had before him. Most of the Chrysorutilia
species from Australia have brilliant yellow heads, but the head of formosa is described as
having a red interfrontal area (‘Frontaux rouges’) and the remainder mainly whitish (‘cdtes
du front et face blanchatres’); the lower part of the face is described as bluish, which appears
to suggest that the epistome is rather metallic. One species that is common in eastern Australia
fits this description very well, as it has the parafrontals and parafacials whitish pollinose (there-
fore contrasting with the red interfrontal area) and often shows distinct metallic colour on the
epistome, and it is therefore from this species that the neotype specimen has been designated
for formosa R.-D. Other characters of the species, including the male genital characters, are
indicated in the accompanying key and text-figure.

It should be noted that Robineau-Desvoidy’s description does not fit splendida (which is a
species with brilliant yellow head), so that even if his use of foymosa and attribution to Donovan
were a lapsus for splendida it is clear that he did not have the true splendida before him. Towns-
end (1938 : 413) cited a ‘Musca formosa [Donovan] (1805)’ but there is no such nominal species
in the works of Donovan; he also stated that formosa R.-D. is a synonym of retusa Fabricius,
but the holotype of the latter still exists and has been examined and found to be completely
distinct from any of the species of Chrysorutilia (belonging in fact in the subgenus Donovanius).
Townsend’s (1915) genus Chrysorutilia rests nomenclaturally upon the statement ‘Genotype,
Rutilia formosa Desvoidy, 1830, Essai Myodaires, 320’ without any account of the characters
defining the taxon; since no characters are cited in this original ‘description’ it is here presumed
that no conflict exists between Townsend’s original meaning of Chrysorutilia and the meaning
given to it in the present work by designation of a neotype for formosa R.-D., the type-species.
It should be recorded, though, that Townsend’s (1938) later meaning of Chrysorutilia (after he
had synonymized formosa with retusa without any confirmatory evidence) is different and his
definition in Manual of Myiology seems to apply better to the concept Donovanius Enderlein,
here regarded as a valid subgenus. There were no specimens cited by Townsend in the original
(1915) proposal of Chrysorutilia, and I hold therefore that Enderlein’s (1936 : 401-408) interpre-
tation of Chrysorutilia Townsend, which is in conformity with the present interpretation based
upon neotype designation for the cited type-species, is taxonomically correct—even if specimens
later held to be formosa by Townsend belong to another concept. (It would not be a case of
misidentified type-species because no specimens were cited in the original description by which
Townsend’s meaning of formosa was established; the type-species of Chrysorutilia must therefore
be the nominal species named by Townsend as type-species, i.e. the formosa R.-D. whose charac-
ters are now pinned down by neotype designation.)

Rutilia imperialis Guérin-Méneville, 1843 : 265.

NEOTYPE¢. AvustrRaia, New South Wales, Mt Wilson, 2.1.1953 (M. F. Day) (in Australian
National Insect Collection, Canberra).

This species was described from a male and a female from ‘Nouvelle-Hollande’ without
further data. It occurs mainly in south-eastern Australia (New South Wales and Victoria)
and as much of the early collecting was done in New South Wales the neotype specimen desig-
nated is from that state.

Rutilia lepida Guérin-Méneville, 1843 : 268.

NEOTYPE ¢g. Ausrratia, Australian Capital Territory, Blundell’s, near Canberra, 19.iv.
1948 (Paramonov) (in Australian National Insect Collection, Canberra).

This species was described from a specimen from ‘Nouvelle-Hollande’ without further data.
The careful description of Guérin-Méneville leaves really no doubt that his name /epida applies
to the species later described by Macquart as Rutilia fulgida. The latter name has not been in
use for any species of Rutilia (and was never, for instance, mentioned by Malloch in his various
papers on Rutiliini), and no disruption of nomenclature arises from fixing the specimen here
cited as neotype of Jepida and sinking Macquart’s name fulgida in new synonymy (see p. 73).
The lectotype of fulgida Macquart (designated by Crosskey, 1971) is in the British Museum

126 R. W. CROSSKEY

(Natural History), and has been directly compared with the specimen chosen as neotype of
lepida; the two primary types are undoubtedly conspecific. Rutilia lepida occurs in south-
eastern Australia from northern New South Wales to Victoria, and the neotype chosen (from
near Canberra) is from a fairly central position in the normal range.

Rutilia pellucens Macquart, 1846 : 305 (177).

NEOTYPE g. AvustTrRaLia, New South Wales, Durras Bay, 15—-30.x.1953 (F. M. Hull) (in
British Museum (Natural History), London).

This species was described from a single specimen (holotype) that stood in Fairmaire’s
collection. Crosskey (1971) considered the type lost, as no Diptera from Fairmaire’s collection
have been located; however, the identity of pellucens is clear from the five specimens of the
species in the British Museum (Natural History) collection that were identified by Macquart
himself. The existence of several species of Rutiiia which are extremely similar to pellucens
and easily confused with it, makes neotype fixation desirable: as there is no data on the specimens
identified by Macquart, other than that they came from Australia, a recently collected specimen
with full data has been chosen as neotype after direct comparison with the specimens seen by
Macquart.

Rutilia regalis Guérin—Méneville, 1831 : plate 21, fig 1.

NEOTYPE g. Avustratia, Australian Capital Territory, Tharwa, 14.ii.1951 (S. J. Para-
monov) (in Australian National Insect Collection, Canberra).

Guérin-Méneville’s excellent coloured plate of this species (published in 1831, earlier than the
text description 1838) shows a large rather uniformly green Rutilia, and has enabled later
workers to recognize vegalis correctly. Specimens of R. rvegalis vary slightly in the intensity
and shade of green colouring; the original plate figure shows a specimen that is rather golden-
green, and the specimen chosen as neotype has a slight golden tinge. The range of the species
is mainly from New South Wales to South Australia.

Rutilia setosa Macquart, 1847 : 94 (78)

NEOTYPE g. Austratia, New South Wales, 4 miles North of Bateman’s Bay, 20.x.1953
(S. J. Paramonov) (in Australian National Insect Collection, Canberra).

Crosskey (1971) showed that the type of this species is lost. It was originally described from
specimens in Bigot’s collection said to be from Tasmania (though there is much uncertainty
about the exact origins of the Australian specimens in Bigot’s collection). Macquart’s descrip-
tion of a brown Rutilia in which the abdomen has the ‘deuxiéme segment muni de huit 4 douze
soies au bord postérieur’ (i.e. T3 with a transverse row of 8-12 median marginal setae in modern
terminology) can only relate to a species of Rutilia s. str. close to vivipara (Fab.); however it is
clearly not vivipara itself, which has entirely reddish yellow legs, because Macquart records
‘pedibus nigris; tibiis testaceis’. In New South Waies there occurs a species close to vivipara
but having the femora and tarsi largely blackish and contrasting with the reddish yellow tibiae,
and it is to this species (until now unnamed) that Macquart’s name sefosa is considered to apply,
and from which the neotype here chosen is designated. Rutilia setosa Macquart differs from
R. vivipara (Fabricius) by having the suprasquamal ridge bare, as well as in the leg coloration
mentioned.

Tachina vivipara Fabricius, 1805 : 309.

NEOTYPE gf. Avsrraria, New South Wales, Barrington Tops, 9.iv.1949 (S. J. Paramonov)
(in Australian National Insect Collection, Canberra).

The provenance of Fabricius’ material has always been assumed to be Australia, though the
locality was cited as ‘Insulis maris pacifici’ in the original description; Fabricius’ description
fits so well with the common eastern Australian Rutilia species that has always been accepted as
vivipara that there is really no doubt about the true identity, and a specimen from New South
Wales fitting with both the past concept of vivipava and with Fabricius’ description is here
designated as neotype. As vivipara is type-species of Rutilia, this neotype is the nomenclatorial

i

REVISIONARY CLASSIFICATION OF RUTILIINI 127

pivot of the Rutiliini. For further information on vivipara see under the discussion of Rutilia

sensu stricto (p. 77).

SUMMARY OF THE) PROPOSED CLASSIFICATION OF THE RUTILIINI

The following synoptic catalogue shows all the genus-group and species-group
taxa of Rutiliini, together with their synonyms, arranged in accordance with the
classification of the tribe here proposed. Names that are considered taxonomically
and nomenclaturally valid are printed in bold-face type, synonyms are indented,
and misidentifications are indicated by brackets around the misapplied names; only
the most important misidentifications are noted. Specific names are alphabetical
within their higher taxon.

Family TACHINIDAE Robineau-Desvoidy, 1830
Subfamily PROSENINAE (Townsend, 1892")
Tribe RUTILIINI Brauer & Bergenstamm, 1889

Genus FORMOSIA Guérin-Méneville, 1843 LACCURA Enderlein, 1936

syn. n

Subgenus FORMOSIA Guérin-Méneville,
1843 S. str.
PANCALA Enderlein, 1936
syn. n.
viridiventris-group
viridiventris Crosskey sp. n.
mirabilis-group
blattina (Enderlein, 1936) comb. n.
bracteata (Enderlein, 1936) comb. n.
callipygos Gerstaecker, 1860
eos (Enderlein, 1936) comb. n.
fervens (Walker, 1861)
flavipennis (Macquart, 1848)
gemmata (Enderlein, 1936) comb. n.
viridescens (Enderlein, 1936)
glorificans (Walker, 1861)
pectoralis (Walker, 1865) syn. n.
fulvipes (Enderlein, 1936)
heinrichiana (Enderlein, 1936)
comb. n.
heinrothi (Enderlein, 1936) comb. n.
mirabilis (Guérin-Méneville, 1831)
plumicornis (Macquart, 1843)
solomonicola Baranov, 1936 stat. n.

Subgenus PSEUDOFORMOSIA Brauer
& Bergenstamm, 1889

excelsa (Walker, 1862) comb. n.
moneta Gerstaecker, 1860
lucigena (Walker, 1861) syn. n.
obscuripennis Brauer & Ber-
genstamm, 1889 (unavail-
able)
pauper (de Meijere, 1904) comb. n.
saturatissima (Walker, 1861)

Subgenus EUAMPHIBOLIA Townsend,
1916
HEGA Enderlein, 1936 syn. n.
CHROMOCHARIS Ender-
lein, 1936 syn. n.
atribasis (Walker, 1861)
complicita (Walker, 1861) comb. n.
pretiosa (Snellen van Vollen-
hoven, 1863) syn. n.

sapphivina (Walker, 1862)
syn. n.

smaragdifera Bigot, 1874 syn.
n.

vividicingens (Enderlein, 1936)
syn. n.

engeli (Enderlein, 1936) comb. n.
faceta (Enderlein, 1936) comb. n.
Susca Crosskey sp. n.

4 See the Introduction for a note on the nomenclatural position of this name.

128 Re Wi

smaragdina Malloch, 1929
speciosa (Erichson, 1842)
fulvipes (Guérin-Méneville,
1843)

Genus FORMODEXIA ¢en. n.
volucelloides (Walker,1861) comb.n.
ignobilis (Walker, 1864) syn.n.
trixoides (Walker, 1861) syn.n.

Genus RUTILODEXIA Townsend, 1915
BOTHROSTIRA Enderlein, 1936 syn. n.
RUTILOSIA Paramonov, 1968 (un-

available)

angustipennis (Walker, 1859)

papua (Bigot, 1880) comb. n.

prisca (Enderlein, 1936) comb. n.
valumensis (Enderlein, 1936)

syn. n.
Genus RUTILIA Robineau-Desvoidy, 1830

Subgenus NEORUTILIA Malloch, 1936
simplex Malloch, 1936

Subgenus AMENIAMIMA subéen. n.
argentifera Bigot, 1874
frontosa (Malloch, 1929) syn.
n.
cingulata (Malloch, 1930)
quadripunctata (Malloch, 1930)
comb. n.

Subgenus CHRYSORUTILIA Townsend,
1915 Stat. n.
PHILIPPOFORMOSIA
Townsend, 1927 syn. n.
HABROTA Enderlein, 1936
ZORAMSCEUS_ Enderlein,

1936 Syn. n.
IDANIA_ Enderlein, 1936
syn. n.

FORMOTILIA Paramonov,
1968 (unavailable)
atrox-group
atrox (Enderlein, 1936) comb. n.

luzona-group
luzona (Enderlein, 1936) comb. n.
townsendi nom. n.
splendida (Townsend, 1927)
(preoccupied)

CROSSKEY

formosa-group

caeruleata (Enderlein, 1936) comb.n.
lineata (Enderlein, 1936)
syn. n.
caesia (Enderlein, 1936) comb. n.
vufibarbis (Enderlein, 1936)
syn. n.
vividescens (Enderlein, 1936)
chersipho (Walker, 1849)
evichsonit Engel, 1925 syn. n.
corona Curran, 1930
cryptica Crosskey sp. n.
decora Guérin-Méneville, 1843
formosa Robineau-Desvoidy, 1830
pubicollis Thomson, 1869
syn. n.
subvittata Malloch, 1929
uzita (Walker, 1849) syn. n.
goerlingiana (Enderlein, 1936)
comb. n.
idesa (Walker, 1849)
imperialis Guérin-Méneville, 1843
vuficornis Bigot, 1880 syn. n.
semtfulva Bigot, 1880 syn. n.
imperialoides Crosskey sp. n.
nana (Enderlein, 1936) comb, n.
panthea (Walker, 1849)
rubriceps Macquart, 1847
angustigena (Enderlein, 1936)
syn. n.
formosina Curran, 1930 Syn. n.
nitens Macquart, 1851 syn. n.
serena (Walker, 1849) syn. n.
splendida (Donovan, 1805)
australasia Gray, 1832
confiuens (Enderlein, 1936)
evanescens (Enderlein, 1936)
transversa Malloch, 1936

Subgenus DONOVANIUS Enderlein,

1936 Stat. n.
PSARONIA Enderlein, 1936
syn. n.
MENEVILLEA Enderlein,
1936 Syn. n.
agalmiodes (Enderlein, 1936)
comb. n.

analoga Macquart, 1851
dubitata Malloch, 1929 syn. n.
bisetosa (Enderlein, 1936) comb. n.
nigrvibasis (Enderlein, 1936)
syn. n.
brunneipennis Crosskey sp. n.
ethoda (Walker, 1849)

REVISIONARY CLASSIFICATION OF RUTILIINI 129

inusta (Wiedemann, 1830)
castanifrons Bigot, 1880syn.n.
castanipes Bigot, 1880 syn. n.
potina (Walker, 1849) syn. n.
spinipectus Thomson, 1869

syn. n.

lepida Guérin-Méneville, 1843
fulgida Macquart, 1846 syn. n.
onoba (Walker, 1849) syn. n.

nigrihirta Malloch, 1935

pellucens Macquart, 1846
imitatoy (Enderlein, 1936)

syn. n.

regalis Guérin-Méneville, 1831
nigva Macquart, 1846 nomen

nudum

retusa (Fabricius, 1775)
aditha (Walker, 1849) syn. n.
vividitestacea Macquart, 1851

syn. n.

sabrata (Walker, 1849)

[tnornata G.-M. sensu auct.
(misident.)]

savaiiensis Malloch, 1935

spinolae Rondani, 1864

transfuga Bezzi, 1928

viridinigra Macquart, 1846
barcha (Walker, 1849) syn. n.
fuscotestacea Macquart, 1846

syn. n.

Subgenus RUTILIA Robineau-Desvoidy,
1830 S. str.
PSARONIELLA Enderlein,
1936 syn. n.
STIRAULAX
1936
confusa (Malloch, 1929)
dentata Crosskey sp. n.
setosa Macquart, 1847
{castanipes Bigot sensu En-
derlein (misident.)]
vivipara (Fabricius, 1805)
desvoidyi Guérin-Méneville,
1843 syn. n.
durvillei Robineau-Desvoidy,
1830 Syn. n.
tnornata Guérin-Méneville,
1843, Syn. n.

Enderlein,

Subgenus GRAPHOLOSTYLUM Mac-
quart, 1851 stat. n.
AGALMIA Enderlein, 1936

syn. n.

albovirida Malloch, 1929
dorsomaculata (Macquart, 1851)

albopicta Thomson, 1869
syn. n.

fuscisquama Malloch, 1930
syn. n.

leucosticta Schiner, 1868
syn. n.

variegata Bigot, 1874 syn. n.
micans Malloch, 1929
subtustomentosa Macquart, 1851

velutina Bigot, 1874 syn. n.

Subgenus MICRORUTILIA Townsend,
1915
PROSENOSTOMA _ Towns-
end, 1932 syn. n.
EUCOMPSA Enderlein, 1936
POGONAGALMIA
Enderlein, 1936, syn. n.
cupreiventris Malloch, 1936 stat. n.
fulviventris Bigot, 1874
flavipes (Brauer & Bergen-
stamm, 1889) syn. n.
hirticeps Malloch, 1929
pallens Curran, 1930 syn. n.
[flavipes B. & B. sensu Towns-
end (misident.)]
liris (Walker, 1849)
media Macquart, 1846
vuficornis (Macquart, 1851)
syn. n.
minor Macquart, 1846
nigriceps Malloch, 1929
nigripes (Enderlein, 1936) comb. n.

Subgenerically unplaced species-group taxa
micropalpis Malloch, 1929
scutellata (Enderlein, 1936) (variety)

Genus AMPHIBOLIA Macquart, 1843

Subgenus AMPHIBOLIA Macquart, 1843
S. SLE.
albocincta (Malloch, 1930)
campbelli Paramonov, 1950
commoni Paramonov, 1968
ignorata Paramonov, 1950
papuana Crosskey sp. n.
valentina Macquart, 1843
vidua (Guérin-Méneville, 1843)

wilsoni Paramonov, 1950

Subgenus PARAMPHIBOLIA Brauer &
Bergenstamm, 1891 stat. n.

130 R. W. CROSSKEY

CHAETOGASTRINA Mal- georgei Malloch, 1929
loch, 1929 Syn. n. minuta Paramonov, 1968
assimilis (Macquart, 1851) comb. n. regina (Malloch, 1936)
stolida (Malloch, 1929) comb. n. testacea (Macquart, 1843)
victoriae (Malloch, 1936)
Genus CHRYSOPASTA Brauer & Bergen- vittata (Macquart, 1855)
stamm, 1889 walkeri Paramonoy, 1968
ROEDERIA Brauer & Bergenstamm,
1893 Genus CHETOGASTER Macquart, 1851
ECHRYSOPASTA Townsend, 1932 CODIUM Enderlein, 1936
elegans (Macquart, 1846) CHAETOGASTER: incorrect subse-
versicoloy Brauer & Bergen- quent spelling
stamm, 1889 argentifera Malloch, 1936
zabirna (Walker, 1849) auriceps Paramonov, 1968
canberrae Paramonov, 1954
Genus PRODIAPHANIA Townsend, 1927 oblonga (Macquart, 1847)
DIAPHANIA Macquart, 1843 (pre- gvatiosa Paramonov, 10954
occupied) syn. n.
[SENOSTOMA sensu auct., not Mac- pellucida Paramonov, 1954
quart] violacea Macquart, 1851
arida Paramonov, 1968 viridis Malloch, 1936
biarmata (Malloch, 1936)
brevitarsis Paramonov, 1968 Nomina nuda (unplaceable)
claripennis Malloch, 1929
commoni Paramonoy, 1968 Diaphania grisea Brauer & Bergenstamm,
cygnus (Malloch, 1936) I89QI
deserta Paramonov, 1968 Formosia viridithovax Bigot, 1874
echinomides (Bigot, 1874) Rutilia accedens Brauer & Bergenstamm,
fullerae Paramonov, 1968 1891
funebris Paramonov, 1968 Rutilia evvonea Paramonov, 1968
furcata (Malloch, 1936) Rutilia humevalis Paramonov, 1968
genitalis Paramonov, 1968 Rutilia incomparabilis Brauer & Bergen-
paratestacea Paramonov, 1968 stamm, 1891
syn. n. Rutilia sovoy Brauer & Bergenstamm, 1891

INDEX-CATALOGUE OF SPECIES-GROUP NAMES AND THEIR TYPES

The index-catalogue given below provides a complete alphabetical list of the
species-group names of Rutiliini, with a summary of essential information on each
primary type. Each species-group name is cited in the original spelling and is
accompanied by the following information:

Author; date and page reference of original publication; genus of original
assignment (in parentheses); status and sex of primary type; authority for
lectotype designation (if relevant) ; data of primary type (when available in the
sequence: locality, altitude, date of collection, name of collector in italics);
type-depository.

Any special annotations, such as explanatory notes on the nomenclatural status
or dating of names, are given after the basic type information (unless the name is
unavailable and therefore has no type). Nomenclaturally available names are
printed in bold type and nomina nuda and other unavailable names are printed in

REVISIONARY CLASSIFICATION OF RUTILIINI 131

non-bold italics. (Surprisingly, there are no instances of primary homonymy in the
Rutiliini.)

A few entries have been enclosed in square brackets and the names printed in
non-bold italics; this indicates, for convenience, those few species that were des-
cribed in Rutiliine genera but which are considered not to belong in the Rutiliini.

The following abbreviations have been used for the museum collections in which
the primary types are housed:

AM Australian Museum, Sydney
AMNH American Museum of Natural History, New York
ANIC Australian National Insect Collection, Canberra

BMNH British Museum (Natural History), London

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels
MHN Musée d’Histoire Naturelle, Lille

MNHN Muséum National d’Histoire Naturelle, Paris

MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin

NM Naturhistorisches Museum, Vienna
NMV National Museum of Victoria, Melbourne
NR Naturhistoriska Riksmuseum, Stockholm

RMNH Rijksmuseum van Natuurlijke Historie, Leiden
SPHTM School of Public Health and Tropical Medicine, Sydney
USNM United States National Museum, Washington D.C.

Full bibliographic references to the works containing the original descriptions of
the species-group taxa, or to the original use of a name if there is no description,
can be found in the ‘References’ section on p. 140.

accedens Brauer & Bergenstamm, 1891: 418 (114) (Rutilia). Nomen nudum (no later validation).

aditha Walker, 1849 : 854 (Dexia). Holotypeg, Austraia (Western Australia, Swan River) :
BMNH, London.

agalmiodes Enderlein, 1936: 412 (Donovanius). Holotype g, AusTRALIA (Queensland,
Cairns, 1907) : MNHU, Berlin.

agalmiodes Enderlein, 1936 : 434 (Prosenosioma, as aberration of ruficorne Macquart, 1851)
Infrasubspecific name without status in nomenclature.

albocincta Malloch, 1930 : 108 (Rutilia). Holotype 9, AUSTRALIA (New South Wales, Barring-
ton Tops, ii.1925, S.U. Zool. Exp.) : ANIC, Canberra.

albopicta Thomson, 1869 : 529 (Rutilia). Holotype 9, AustrRaLta (New South Wales, Sydney,
Kinb.) : NR, Stockholm.

albovirida Malloch, 1929 : 307 (Rutilia (Senostoma)). Holotype 9, AUSTRALIA (Queensland,
Yeppoon, x.1924, A. Musgrave) : AM, Sydney.

analoga Macquart, 1851 : 191 (218) (Rutilia). Holotype 2 [publ. as 3], AusTRALIA (? New
South Wales [publ. as ‘Tasmanie’]) : MNHN, Paris.

[angustecarinata Macquart, 1848 : 211 (51) (Rutilia). Not Rutiliini. g holotype discovered
in 1971 in the remnants of Payen’s collection, formerly in the Municipal Museum at Tournai
and now in IRSNB, Brussels, shows that this nominal species is not a Rutiliine but belongs
in the Tachinid tribe Nemoraeini.]

angustigena Enderlein, 1936: 403 (Chrysorutilia). Lectotype 3, by present designation
(p. 117), AUSTRALIA (Queensland, Herberton, i.t910) : MNHU, Berlin.

132 Re We GCROSSKENW

angustipennis Walker, 1859 : 101 (Rutilia). Holotype g, Inpones1a (Aru Islands, A. R.
Wallace) : BMNH, London.

argentifera Bigot, 1874 : 464 (Rutilia). Lectotype g, by designation of Crosskey (1971 : 300),
AUSTRALIA (New South Wales, Sydney) : BMNH, London.

argentifera Malloch, 1936 : 19 (Chaetogaster). Holotype 2, AUSTRALIA (Victoria, Gisborne,
25.11.1923, G. Lyell) : SPHTM, Sydney.

arida Paramonov, 1968 : 397 (Prodiaphania). Holotype, AusTRALIA (Victoria, Little Desert,
5 miles S. of Kiata, 13.ii.1956, J. F. B. Common) : ANIC, Canberra.

assimilis Macquart, 1851 : 192 (219) (Rutilia). Lectotype g, by designation of Crosskey
(1971 : 285), AUSTRALIA (Tasmania) : MNHN, Paris.

Macquart cited the type-locality as ‘Nouvelle-Hollande: céte orientale’ in the original
description, but Crosskey (1970) established, from accession reference numbers on the
type-material, that Tasmania is the correct provenance.

atribasis Walker, 1861b : 288 (Rutilia). Lectotype 2, by present designation (p. 119),
InpDonEs1A (Moluccas, Batjan, A. R. Wallace) : BMNH, London.

atrox Enderlein, 1936: 408 (Idania). Holotype 9, PHILIPPINE REPUBLIC (Luzon, Imugan,
6.vi.1918, Georg Bottcher) : MNHU, Berlin.

auriceps Paramonov, 1968 : 371 (Chaetogaster). Holotype 9, AUSTRALIA (Queensland,
Collinsville, 15.1x.1950, E. F. Riek) : ANIC, Canberra.

australasia Gray in Cuvier, 1832: pl. 114, fig. 1 & p. 793 [index] (Rutilia). Holotype or
syntypes [2], AUSTRALIA : lost.

The description of this nominal species consists of a good figure by Gray clearly showing
a 2 Rutilia, and of the words ‘Black and blue’ appearing in the index on p. 793 of the
second Insecta volume of Cuvier’s The Animal Kingdom. No specimen or specimens that
could have formed the basis for Gray’s figure have been located, and the type-material is
therefore presumed lost. Walker (1849 : 863) synonymized the name R. austvalasia Gray
with R. splendida (Donovan); this synonymy, which is fully justified by Gray’s excellent
figure, is here upheld as certainly correct.

barcha Walker, 1849 : 857 (Dexia). Holotype 9, AUSTRALIA : BMNH, London.

biarmata Malloch, 1936 : 14 (Senostoma). Holotype g, AustRatia (South Australia, J. B.
Cleland) : SPHTM, Sydney.

bisetosa Enderlein, 1936: 414 (Psaronia). Lectotype 2, by present designation (p. 119),
AUSTRALIA (Western Australia, Wurarga, Marloo Station, 1935, Goerling) : MNHU,
Berlin.

blattina Enderlein, 1936 : 423 (Pancala). Holotype 9, INDoNnEsIA (Celebes, Latimodjong
Mts, 1300-1500 m, vii.1930, G. Heinrich) : MNHU, Berlin.

bracteata Enderlein, 1936 : 425 (Pancala). Holotype g, Papua New Guinea (southern New
Ireland [‘Siid-Neu-Mecklenburg’], exped. Friederici) : MNHU, Berlin.

brevitarsis Paramonov, 1968 : 395 (Prodiaphania). Holotype g, AUSTRALIA (New South
Wales, near Queanbeyan, 10.ii1.1953, S. J. Pavamonov) : ANIC, Canberra.

brunneipennis Crosskey, 1972 : (present paper) (Rutilia (Donovanius)). Holotype 2, SoLo-
MON IsLanps (Guadalcanal, Suta, 27.vi.1956, E. S. Brown) : BMNH, London.

caeruleata Enderlein, 1936 : 402 (Chrysorutilia). Lectotyped, by present designation (p. 117),
AUSTRALIA (Western Australia, Wurarga, Marloo station, ix.1935, A. Goerling) : MNHU,
Berlin.

caesia Enderlein, 1936 : 402 (Chrysorutilia). Lectotype g, by present designation (p. 117),
AUSTRALIA (Clement) : MNHU, Berlin.

callipygos Gerstaecker, 1860 : 198 (Formosia). Holotype 2 New Guinea (Felder) : MNHU,
Berlin.

campbelli Paramonov, 1950 : 523 (Amphibolia). Holotype g, AusTRALIA (Australian Capital
Territory, Blundell’s near Canberra, 23.xi1.1938, T. G. Campbell) : ANIC, Canberra.

canberrae Paramonov, 1954 : 277 (Chaetogaster). Holotype §, AUSTRALIA (Australian Capital
Territory, Canberra, Black Mt, 30.xii.1929, G. A. Currie) : ANIC, Canberra.

castanifrons Bigot, 1880 : 88 (Rutiia). Holotype 2, AusTRALIA : BMNH, London.

REVISIONARY CLASSIFICATION OF RUTILIINI 133

castanipes Bigot, 1880 : 87 (Rutilia). Lectotype 2, by designation of Crosskey (1971 : 300),
AUSTRALIA : BMNH, London.

chersipho Walker, 1849 : 864 (Dexia). Neotyped, by present designation (p. 123), AUSTRALIA
(Western Australia, Waroona, 21.xi.1908, G. F. Berthoud) : BMNH, London.

cingulata Malloch, 1930: 105 (Formosia). Holotype 3, Austratia (New South Wales,
Wentworth Falls, 14.xii.1923, Harrison) : ANIC, Canberra.

claripennis Malloch, 1929 : 292 (Prodiaphania, as variety of testacea Macquart, 1843). Holo-
type J, AUSTRALIA (Western Australia, King George’s Sound) : AM, Sydney.

commoni Paramonov, 1968 : 363 (Amphibolia). Holotype 2, AustrRaria (Victoria, Gram-
pians, Fyan’s Creek, 11.ii.1956, J. F. B. Common) : ANIC, Canberra.

commoni Paramonov, 1968 : 389 (Prodiaphania). Holotype g, AUSTRALIA (Victoria, Little
Desert, 5 m. S. of Kiata, I. F. B. Common) : ANIC, Canberra.

complicita Walker, 1861b : 288 (Rutilia). Holotype g [not 9], InponeEsiA (Moluccas, Batjan,
A. R. Wallace) : BMNH, London.

confluens Enderlein, 1936: 407 (Chrysorutilia, as variety of splendida Donovan, 1805)
Lectotype 3, by present designation (p. 118), AUSTRALIA (Queensland) : MNHU, Berlin.

confusa Malloch, 1929 : 309 (Formosia). Holotype g, Austratia (South Australia, Kangaroo
Island, deep creek 20 m. from Kingscote, E. Troughton) : AM, Sydney.

corona Curran, 1930 : 3 (Rutilia). Holotype g, AustRALIA (New South Wales, H. Edwards) :
AMNH, New York.

cryptica Crosskey, 1972 : (present paper) (Rutilia (Chrysorutilia)). AUSTRALIA (South Austra-
lia, near Moonta, 16.xi.1904, W. Wesche) : BMNH, London.

cupreiventris Malloch, 1936: 18 (Rutilia (Microrutilia) as variety of ruficornis Macquart,
1851). Holotype g, AustraLt1a (New South Wales, Barrington Tops, ii.1925, S.U. Zool.
Exp.) : SPHTM, Sydney.

cygnus Malloch, 1936: 15 (Senostoma). Holotype g, AustRaALia (Western Australia, Swan
River, L. J. Newman) : SPHTM, Sydney.

decora Guérin-Méneville, 1843 : 266 (Rutilia). Neotype g, by present designation (p. 124),
AUSTRALIA (Queensland, Burpengary, T. L. Bancroft) : BMNH, London.

dentata Crosskey, 1972 : (present paper) (Rutilia (Rutilia)). Holotype g, AUSTRALIA (Victoria,
Monbulk) : BMNH, London.

deserta Paramonov, 1968 : 398 (Prodiaphania). Holotype g, AusTRALIA (New South Wales,
Wanaaring, 29.x.1949, S. J. Paramonov) : ANIC, Canberra.

desvoidyi Guérin-Méneville, 1843 : 269 (Rutilia). Lectotype 2, by present designation
(p. 119), AUSTRALIA : MNHN, Paris.

[diversa Paramonov, 1954 : 280 (Chaetogaster). Not Rutiliini as herein defined.]

dorsomaculatum Macquart, 1851 : 196 (223) (Grapholostylum). Lectotype g, by designation
of Crosskey (1971 : 271), AUSTRALIA (‘Tasmanie’, probably error for New South Wales) :
MNHN, Paris.

[dubia Macquart, 1846 : 311 (183) (Rutilia). Not Tachinidae, see Crosskey (1971 : 285).]

dubitata Malloch, 1929 : 303 (Rutilia). Holotype 9, AusTRALIA (New South Wales, Jindabyne,
11.1889, Helms) : AM, Sydney.

durvillei Robineau-Desvoidy, 1830 : 321 (Rutilia). Neotyped, by present designation (p. 124),
AUSTRALIA (New South Wales, Hawkesbury River, 28.xi.1914) : BMNH, London.

echinomides Bigot, 1874 : 466 (Rutilia). Holotype 9, AUSTRALIA : BMNH, London.

elegans Macquart, 1846 : 309 (181) (Rutilia). Holotype g, AusTRaLia (New South Wales,

Sydney) : BMNH, London.

engeli Enderlein, 1936 : 431 (Laccura). Lectotype 2, by present designation (p. 118), INDo-
NESIA (Banda Islands, Run [= Roon] Island, south of Ceram) : MNHU, Berlin.

eos Enderlein, 1936 : 423 (Pancala). Lectotype 3, by present designation (p. 118), INDONESIA
(Celebes, Bonthain, Wawa Karaeng, 1100 m., viii.1931, G. Heinrich) : MNHU, Berlin.

evichsonit Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum (later validated
as evichsonii Engel, q.v.).

erichsonii Engel, 1925 : 363 (Rutilia). Lectotype 2, by fixation of Malloch (1929 : 297),

134 Re W. CROSSKEY

AUSTRALIA (Western Australia, Swan River) : NM, Vienna.
evvonea Paramonov, 1968 : 356, 361 (Rutilia). Nomen nudum.

This name was published in Paramonov’s (1968) posthumous paper on Rutiliini in the
discussion of the genus Amphibolia Macquart and in the preamble to the description there
given of A. albocincta (Malloch); there is no definition of the taxon, and the name erronea
is unavailable (nomen nudum).

ethoda Walker, 1849 : 856 (Dexia). Holotype 2, AUSTRALIA (Western Australia, Swan River) :
BMNH, London.

evanescens Enderlein, 1936: 407 (Chrysorutilia, as variety of splendida Donovan, 1805).
Holotype 3, AusTRALIA (Damel) : MNHU, Berlin.

The holotype bears a label reading ‘N. Holl/sept. Damel’ and Enderlein (1936 : 407)
interpreted this as northern Australia (“Nord-Australien’); this seems doubtful, and the
provenance within Australia is here considered unknown.

excelsa Walker, 1862: 19 (Rutilia). Holotype 3, InponEsIA (Moluccas, Ternate, A. R.
Wallace) : BMNH, London.

faceta Enderlein, 1936: 422 (Hega). Holotype g, Austratia (N. Queensland) : MNHU,
Berlin.

fervens Walker, 1861 : 288 (Rutilia). Holotype 2, InponEs1a (Moluccas, Batjan, A. R.
Wallace) : BMNH, London.

flavipennis Macquart, 1848: 210 (50) (Rutilia). Holotype g, Java (Payen) : IRSNB
Brussels (ex Municipal Mus., Tournai).

Crosskey (1971 : 286) could not locate the original material of this species during work
on Macquart’s types, but the gf holotype was located during the present work in the rem-
nants of Payen’s collection (now in Brussels) from the Municipal Museum, Tournai. It
consists of the eaten out shell of head and thorax and abdominal venter, but all legs and
genitalia are present and the identity is clear; the body is discoloured.

flavipes Brauer & Bergenstamm, 1889: 126 (58) (Senostoma). Lectotype 2, by present
designation (p. 121), AUSTRALIA (‘New Holland’) : NM, Vienna.

formosa Robineau-Desvoidy, 1830 : 320 (Rutilia). Neotypedg, by present designation (p. 124)
AUSTRALIA (New South Wales, near Lake George, 25.xi.1953, S. J. Paramonov) : ANIC,
Canberra.

formosina Curran, 1930 : 2 (Rutilia). Holotype g, Austratia : AMNH, New York.

frontosa Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum (later validated
as frontosa Malloch, q.v.).

frontosa Malloch, 1929: 310 (Formosia). Holotype g, AusTRALIA (New South Wales,
Jindabyne, 3000 ft, iii.1889, Helms) : AM, Sydney.

fu’gida Macquart, 1846 : 308 (180) (Rutilia). Lectotype g, by designation of Crosskey (1971 :
286), AUSTRALIA (New South Wales, Sydney) : BMNH, London.

fullerae Paramonov, 1968 : 393 (Prodiaphania). Holotype 3, AUSTRALIA (New South Wales,
Barrington Tops, 29-30.xii.1934, M. E. Fullev) : ANIC, Canberra

fulvipes Enderlein, 1936 : 426 (Pancala, as variety of glorificans Walker, 1861) Holotype Q,
Papua NEw Guinea (Milne Bay, Micholitz) : MNHU, Berlin.

fulvipes Guérin-Méneville, 1843 : 273 (Rutilia). Holotype 9, AUSTRALIA : lost.

fulviventris Bigot, 1874 : 465 (Rutilia). Lectotype 9, by designation of Crosskey (1971 : 301),
AUSTRALIA (Tasmania) : BMNH, London.

funebris Paramonov, 1968 : 391 (Prvodiaphania) Holotype g, AusTRALIA (South Australia,
4o m. S.W. of Iron Knob, 23.x.1958, J. F. B. Common) : ANIC, Canberra.

furcata Malloch, 1936 : 14 (Senostoma). Holotype g, AusTRAtiA (Australian Capital Territory
Canberra, 16.xii.1928, MM. Fullev) : ANIC, Canberra.

fusca Crosskey, 1972 : (present paper) (Formosia (Euamphibolia)). Holotype ¢, INDONESIA
(Moluccas, Seram, Mansela, 2500 ft, 1919, Pratt) : BMNH, London.

fuscisquama Malloch, 1930 : 107 (Rutilia, as variety of leucosticta Schiner, 1868). Holotype 9,
AUSTRALIA (New South Wales, Barrington Tops, Allyn Range, 11.1925, on Leptospermum,
S.U. Zool. Exp.) : ANIC, Canberra.

REVISIONARY CLASSIFICATION OF RUTILIINI 135

fuscotestacea Macquart, 1846 : 306 (178) (Rutilia). Holotype 9, AusTRALIA (New South
Wales, Sydney) : BMNH, London.

gemmata Enderlein, 1936 : 424 (Pancala). Lectotype g, by present designation (p. 118),
Papua NEw Guinea (New Britain, Ralum, Karakaul, 30.viii.1896, in high forest, F. Dahl) :
MNHU, Berlin.

genitalis Paramonov, 1968: 400 (Prodiaphania). UHolotype g, AUSTRALIA (Queensland,
Herberton, ii.1911) : BMNH, London.

georgei Malloch, 1929 : 292 (Prodiaphania). Holotype g, AustTRaLia (Western Australia,
King George’s Sound) : AM, Sydney.

glorificans Walker, 1861a : 241 (Rutilia). Holotype g, INDONESIAN NEw GUINEA (Dorey,
A. R. Wallace) : BMNH, London.

goerlingiana Enderlein, 1936: 404 (Chrysorutilia). Lectotype 9, by present designation
(p. 117), AUSTRALIA (Western Australia, Wurarga, Marloo Station, i.1935, A. Goerling) :
MNHU, Berlin.

gratiosa Paramonoy, 1954 : 283 (Chaetogaster). Holotype g, AUSTRALIA (Victoria, Wonnan-
gatta River, Gibraltar Point, 12.xii.1949, T. G. Campbell) : ANIC, Canberra.

grisea Brauer & Bergenstamm, 1891 : 417 (113) (Diaphania). Nomen nudum (no later valida-
tion).

heinrichiana Enderlein, 1936 : 426 (Pancala). Holotype 3, INpones1a (Celebes, Bonthain,
Wawa Karaeng, 1100 m, ix—x.1931 G. Heinrich) : MNHU, Berlin.

heinrothi Enderlein, 1936 : 425 (Pancala). Holotype 9, Papua NEw Guinea (New Britain,
Matupi, xii.1900-v.1901, Heinroth) : MNHU, Berlin.

hirticeps Malloch, 1929 : 305 (Rutilia (Senostoma)). Holotype g, AusTRALIA (New South
Wales, Monaro, Moonbar, 3000-3500 ft, iii.1889, Helms) : AM, Sydney.

humeralis Paramonov, 1968 : 355 (Rutilia). Nomen nudum.

This name was published in the expression ‘Rutilia humeralis group’ in Paramonov’s
(1968) posthumous paper on Rutiliini in the key to genera (p 355); there is no definition
of the specific taxon and the name is unavailable (nomen nudum).

idesa Walker, 1849 : 858 (Dexia). Holotype ¢ [not 9], Austratta : BMNH, London.
ignobilis Walker, 1864 : 238 (Rutilia). Holotype 2, InponeEsia (Moluccas, Halmahera, A. R.
Wallace) : BMNH, London

Walker, in the original publication, cited the provenance of this nominal species as
Gorrite (a locality in Brazil), but the specimen was actually collected by Wallace in the
island of Gilolo (= Halmahera) and is-so labelled

ignorata Paramonov, 1950 : 522 (Amphibolia) Holotype g, AustraLia (New South Wales
Tindery 1.i.1938, Mackerras) : ANIC, Canberra.

imitator Enderlein, 1936: 412 (Donovanius). Holotype 3g, Austratia (Behy) : MNHU,
Berlin.

imperialis Guérin-Méneville, 1843 : 265 (Rutilia). Neotyped, by present designation (p. 125),
AUSTRALIA (New South Wales, Mt Wilson, 2.i1.1953, M. F. Day) : ANIC, Canberra.

imperialoides Crosskey, 1972 : (present paper) (Rutilia (Chrysorutilia)). Holotypeg, AusTRA-
LIA (New South Wales, Wee Jasper, xii.1920) : BMNH, London.

imcomparabilis Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum (no later
validation).

inornata Guérin-Méneville, 1843 : 268 (Rutilia). Lectotype 2, by present designation (p. 119),
AUSTRALIA : MNHN, Paris.

inusta Wiedemann, 1830 : 306 (Zachina). Lectotype 2, by present designation (p. 121),
AUSTRALIA (New South Wales, Sydney [‘Port Jackson’], Eschscholtz) : MNHU, Berlin.

lepida Guérin-Méneville, 1843 : 268 (Rutilia). Neotype g, by present designation (p. 125),
AustTRALtia (Australian Capital Territory, Blundell’s near Canberra, 19.iv.1948, Pavamonov) :
ANIC, Canberra.

leucosticta Schiner, 1868 : 319 (Rutilia). Holotype 2, AusTratia : NM, Vienna.

Schiner, in the original publication, cited the provenance of this species as New Zealand
(‘Neuseeland’), but the holotype is labelled as from Australia (‘New Holland’); the latter

136 R. W. CROSSKEY

is certainly correct, as the Rutiliini are absent from New Zealand. Engel (1925 : 364)
and Malloch (1928 : 659) have correctly noted the Australian provenance.

lineata Enderlein, 1936 : 407 (Chrysorutilia). Lectotype g, by present designation (p. 118),
AUSTRALIA (Western Australia, Wurarga, Marloo Station, iv.1935, A. Goervling) : MNHU,
Berlin.

liris Walker, 1849 : 882 (Musca). Holotype ?, AUSTRALIA (? Tasmania) : BMNH, London.

The type-locality of livis is unknown, but the species to which the holotype belongs is

found in Tasmania, and this is the most probable provenance of the type.

lucigena Walker, 1861a: 242 (Rutilia). Lectotype g, by present designation (p. 120),
INDONESIAN NEw GuINEA (Dorey, A. R. Wallace) : BMNH, London.

luzona Enderlein, 1936: 406 (Chrysorutilia). Wolotype g, PHILIPPINE REPUBLIC (Luzon,
Imugan, 30.vi.1918, Georg Boettcher) : MNHU, Berlin.

media Macquart, 1846 : 310 (182) (Rutilia). Lectotype g, by designation of Crosskey (1971 :
286), AUSTRALIA (Tasmania) : MNHN, Paris.

micans Malloch, 1929 : 299 (Rutilia). Holotype 9, AusTRALIA (New South Wales, Kosciusko,
5000 ft, iii.1889, Helms) : AM, Sydney.

micropalpis Malloch, 1929 : 298 (Rutilia). Holotype, 9 AustraL1a (New South Wales, Como
near Sydney, 7.xi.1923, J. T. Campbell & A. Musgrave) : AM, Sydney.

minor Macquart, 1846 : 310 (182) (Rutilia). Lectotype g, by designation of Crosskey (1971 :
286), AUSTRALIA (Tasmania) : MNHN, Paris.

minuta Paramonov 1968 : 399 (Prodiaphania). Holotype g AUSTRALIA (Queensland,
Gordonvale, 1923) : ANIC, Canberra.

mirabilis Guérin-Méneville, 1831 : plate 21, fig. 2 (Rutilia); 1838 : 2906 (Musca). Holotype 9
(head lost), INDONESIAN NEW GuINEA (Fak-Fak [‘Offak’]) : MNHN, Paris.

This name is available from publication of the plate figure (as Rutilia mirabilis) in 1831,

which pre-dated the text description (as Musca mirabilis) published in 1838.

moneta Gerstaecker, 1860 : 200 (Formosia). Holotype 92, NEw Guinea (Felder) : MNHU,
Berlin.

nana Enderlein, 1936 : 404 (Chrysorutilia). Holotype 2, INDoNEsIA (Kai [= Kei] Islands) :
MNHU, Berlin.

nigva Macquart, 1846 : 305 (177) (Rutilia). Nomen nudum (no later validation).

Specific named cited without definition of the taxon, but not as a synonym, in the

original description of Rutilia pellucens Macquart, and unavailable (nomen nudum).

nigribasis Enderlein, 1936 : 411 (Donovanius, as variety of fulgidus Macquart, 1846). Lecto-
type g, by present designation (p. 118). AUSTRALIA (Western Australia Wurarga, Marloo
Station, 6.vi.1935, A. Goerling) : MNHU, Berlin.

nigriceps Malloch, 1929 : 306 (Rutilia (Senostoma)). Holotype g, AUSTRALIA (New South
Wales, East Dorrigo, Ulong, W. Heron) : AM, Sydney.

nigrihirta Malloch, 1935 : 349 (Rutila (Rutilia)). Holotype g, Samoa (Upolu, Malololelei,
2000 ft, 2.vili.1925, Buxton & Hopkins) : BMNH, London.

nigripes Enderlein, 1936 : 435 (Prosenostoma). Lectotype g, by present designation (p. 119),
AUSTRALIA (Queensland, Herberton, 3700 ft, ii.1911, Dodd) : MNHU, Berlin.

[nigrithovax Macquart, 1851 : 190 (217) (Rutilia). Not Rutiliini as herein defined.]

nitens Macquart, 1851 : 189 (216) (Rutilia). Holotype 2, Inp1a : MNHN, Paris.

oblonga Macquart, 1847 : 92 (76) (Rutilia). Holotype g, AUSTRALIA : BMNH, London.

obscuripennis Brauer & Bergenstamm, 1889 : 126 (58) (Formosia); Brauer & Bergenstamm,
1891 : 434 (130) (Pseudoformosia) & 435 (131) (Formosia).

Name attributed to Bigot but published by Brauer & Bergenstamm in synonymy with
Pseudoformosia moneta (Gerstaecker) and unavailable under Article 11 (d) of the Inter-
national Code of Zoological Nomenclature, 1961.

onoba Walker, 1849 : 859 (Dexia). Holotype fg, AUSTRALIA : BMNH, London.
pallens Curran, 1930: 2 (Rutilia). Holotype g, AustRat1a (New South Wales) : AMNH,
New York.

REVISIONARY CLASSIFICATION OF RUTILIINI 137

panthea Walker 1849 : 862 (Dexia). Holotype 9 AusTRALtIaA (Western Australia) : BMNH,
London.

papua Bigot, 1880 : 87 (Formosia). Lectotype ? [not g] by designation of Crosskey (1971 :
298), NEw Guinea (L. Laglaise) : BMNH, London.

papuana Crosskey, 1972 : (present paper) (Amphibolia (Amphibolia)). Holotype 9, NEw
GuINngEA (Murmur Pass, 8600 ft, x.1961, W. W. Brandt) : BMNH, London.

paratestacea Paramonov, 1968 : 397 (Prodiaphania). Holotype 3, AusTRALIA (New South
Wales, Wee Jasper, 26.i1.1936, M. Fuller) ; ANIC, Canberra.

pauper de Meijere, 1904 : 178 (Pseudoformosia). Lectotype g, by present designation (p. 119),
InponeEsIA (Moluccas, N. Halmahera, Bernstein) : RMNH, Leiden.

pectoralis Walker, 1865 : 114 (Rutilia). Holotype 9, NEw Guinea (A. R. Wallace) : BMNH,
London.

pellucens Macquart, 1846 : 305 (177) (Rutilia). Neotype 3, by present designation (p. 126),
AustTRALiA (New South Wales, Durras Bay, 15—30.x.1953, F. M. Hull) : BMNH, London.

pellucida Paramonov, 1954 : 276 (Chaetogaster, as variety of argentifera Malloch, 1936).
Holotype gj, AustTRALIA (New South Wales, Toronto) : ANIC, Canberra.

plumicornis Macquart, 1843 : 239 (82) (Rutilia). INDONESIAN NEW GuINEA (Fak-Fak
[‘Offak’]) : type-material lost or possibly represented by the holotype of Rutilia mirabilis
Guérin-Méneville.

It is possible that Macquart’s name plumicornis, which he attributed to Guérin-Méneville
in the original description, is based upon one and the same type-specimen as Rutilia
mirabilis Guérin-Méneville: for further discussion of this possibility see Crosskey (1971 :
280).

potina Walker, 1849 : 857 (Dexia). Holotype 9, AUSTRALIA (Tasmania) : BMNH, London.
pretiosa Snellen van Vollenhoven, 1863 : 15 (Rutilia) Lectotype g, by present designation
(p. 120), INDonEsIA (Moluccas, Ternate, Bernstein) : RMNH, Leiden.
_ prisca Enderlein, 1936 : 413 (Bothvostiva). Holotype 3, Papua NEw Guinea (New Britain,
| Kinigunang [publ. as ‘Kinikunang’], C. Ribbe) : MNHU, Berlin.
_ pubicollis Thomson, 1869 : 530 (Rutilia). Lectotype g, by present designation (p. 120),
AUSTRALIA (New South Wales, Sydney, Kinb.) : NR, Stockholm.
quadripunctata Malloch, 1930: 104 (Formosia). Holotype 3g, AUSTRALIA (Queensland,
Eidsvold, xii.1922, Mackerras) : ANIC, Canberra.
ralumensis Enderlein, 1936 : 409 (Idania). Lectotype 2, by present designation (p. 118),
| Papua NEw Guinea (New Britain, Ralum, 30.viii.1896, F. Dahl) : MNHU, Berlin.
| regalis Guérin-Méneville, 1831 : plate 21, fig. 1 (Rutilia); 1838 : 295 (Musca). Neotype ¢
by present designation (p. 126), AustTRatia (Australian Capital Territory, Tharwa
14.11.1951, S. J. Paramonov) : ANIC, Canberra.
This name is available from publication of the plate figure (as Rutilia regalis) in 1831,
which pre-dated the text description (as Musca regalis) published in 1838.
regina Malloch, 1936 : 14 (Senostoma). Holotype g, AUSTRALIA (Queensland, Eidsvold, xii.
1922) : SPHTM, Sydney.
retusa Fabricius, 1775 : 775 (Musca). Holotype 9, AustRaLIA : BMNH, London (Sir Joseph
Banks coll.).
rubriceps Macquart, 1847 : 92 (76) (Rutilia). Holotype 9, AUSTRALIA (‘Tasmanie’, possibly in
error) : BMNH, London.
This species was described from Tasmania, but the true provenance of the holotype is
more probably Queensland (Crosskey, 1971 : 289).
rufibarbis Enderlein, 1936 : 405 (Chrysorutilia). Lectotype g, by present designation (p. 118),
AUSTRALIA (Western Australia, Wurarga, Marloo Station, ix—x.1934, A. Goerling) : MNHU,
Berlin.
uficornis Bigot, 1880 : 88 (Rutilia). Holotype fg, AustRaLiA : BMNH, London.
This name is a junior secondary homonym in Rutilia of R. ruficornis (Macquart, 1851) ;
no new name is required as ruficornis Bigot is a synonym of R. imperialis Guérin—Méneville.

citar sataneee

138 R. W. CROSSKEY

ruficornis Macquart, 1851 : 193 (220) (Diaphania). Holotype g, AusTRALIA (Tasmania) :
MNHN, Paris.

sabrata Walker, 1849 : 855 (Dexia). Holotype 2, AUSTRALIA : BMNH, London.

sapphirina Walker, 1862 : 9 (Rutilia). Holotype g, InponrEs1A (Moluccas, Halmahera [as
‘Gilolo’], A. R. Wallace) : BMNH, London.

saturatissima Walker, 1861b : 287 (Rutilia). Lectotype J, by present designation (p. 120),
InponeEsia (Moluccas, Batjan, A. R. Wallace) : BMNH, London.

savaiiensis Malloch, 1935 : 350 (Rutilia (Rutilia)). Wolotype g, Samoa (Savaii Island,
Fagamalo, xi.1925, Buxton & Hopkins) : BMNH, London.

scutellata Enderlein, 1936 : 405 (Chrysorutilia, as variety of media Macquart, 1846). Holo-
type 9, AUSTRALIA (South Australia, Adelaide, Schomburgk) : MNHU, Berlin.

semifulva Bigot, 1880 : 89 (Rutilia). Lectotype J, by designation of Crosskey (1971 : 301),
AUSTRALIA : BMNH, London.

serena Walker, 1849 : 865 (Dexia). Neotype 9, by present designation (p. 123), Inp1a (Maha-
rashtra, Purandhar, near Poona, c. 3000 ft, on bush, E. P. Sewell) : BMNH, London.

setosa Macquart, 1847: 94 (78) (Rutilia). Neotype g, by present designation (p. 126),
AUSTRALIA (New South Wales, 4 m. N. of Bateman’s Bay, 20.x.1953, S. J. Pavamonov) :
ANIC, Canberra.

simplex Malloch, 1936: 17 (Rutilia (Neorutilia)). Holotype g, AUSTRALIA (Queensland,
Eidsvold, xii.1922) : SPHTM, Sydney.

smaragdifera Bigot, 1874 : 462 (Formosia). Lectotype g, by designation of Crosskey (1971 :
298), INDoNnEsIa (Moluccas, Batjan) : BMNH, London.

smaragdina Malloch, 1929 : 312 (Formosia). Holotype 9, AUSTRALIA (N. Queensland,
Gordonvale, 2.x.1917, E. Jarvis) : AM, Sydney.

solomonicola Baranov, 1936: 101 (Formosia, as subspecies of mirabilis Guérin-Méneville,
1831). Lectotype g, by designation of Sabrosky & Crosskey (1969 : 44), SoLomoNn IsLANDS
(Guadalcanal, Kaukau [publ. as ‘Kankau’], 21.vili.1934, R. J. A. W. Lever) : BMNH,
London.

sovoy Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum (no later validation).

speciosa Erichson, 1842 : 273 (Rutilia). Lectotype 9, by present designation (p. 120),
AUSTRALIA (Tasmania, Schayer) : MNHU, Berlin.

spinipectus Thomson, 1869 : 530 (Rutilia). Lectotype g, by present designation (p. 120),
AUSTRALIA (New South Wales, Sydney, Kinb.) : NR, Stockholm.

spinolae Rondani, 1864 : 23 (Rutilia). Holotype or syntypes, sex unknown, AUSTRALIA : not
located, probably lost.

The introduction to the paper in which this name was published suggests that the type-
material should be in Naples, but (in spite of careful search of museum collections at
Naples and Portici recently made on request) it has not been located there or elsewhere in
Italy. The original material is probably lost, and the name remains enigmatic.

splendida Donovan, 1805: plate fig. (unnumbered) and description (unpaginated) (Musca).
Neotype 3, by present designation (p. 123), AUSTRALIA (Queensland, Brisbane, 8.ix.1907,
A.J. C. Wightman) : BMNH, London.

splendida Townsend, 1927 : 282 (Philippofoymosia). Holotype 9, PHILIPPINE REPUBLIC
(Nueva Viscaya. Imugin, Baker) : USNM, Washington D.C.

This name is a junior secondary homonym in Rutilia of R. sblendida (Donovan, 1805),

see townsendi nom. n.

stolida Malloch, 1929 : 313 (Chaetogastvina). Holotype g, AUSTRALIA (New South Wales,
Barrington Tops, 20.1.1927, T. G. Campbell) : AM, Sydney.

subtustomentosa Macquart, 1851 : 191 (218) (Rutilia). Holotype g, AUSTRALIA (Tasmania) :
MNHN, Paris.

subvittata Malloch, 1929 : 295 (Rutilia, as variety of formosa Robineau-Desvoidy, 1830).
Holotype 3, AUSTRALIA (Western Australia, King George’s Sound) : AM, Sydney.

testacea Macquart, 1843 : 278 (121) (Diaphania). Holotype or syntypes g, AUSTRALIA :
MHN, Lille.

REVISIONARY CLASSIFICATION OF RUTILIINI 139

For further detail on original material in Macquart’s collection in Lille see Crosskey

(1971: 267).

townsendi nom. n. for Philippoformosia splendida Townsend, 1927, junior secondary homonym
preoccupied in Rutilia by R. splendida (Donovan, 1805). For type-data see under splendida
Townsend above.

transfuga Bezzi, 1928 : 192 (Rutilia). Holotype 2, Fiji (Viwa, 26.x.1921, H. W. Simmonds) :
BMNH, London.

Bezzi, in the original publication, recorded the type-locality in Fiji as ‘Suva’, but this
was clearly an error due to misreading of the handwritten word ‘Viwa’ given on the data
label of the holotype. Viwa, a small island of the Fiji group lying north-west of Viti
Levu, is the correct type-locality.

transversa Malloch, 1936: 15 (Rutilia). Holotype g, AUSTRALIA (Western Australia, Swan
River, J. Clavk) : SPHTM, Sydney.

trixoides Walker, 1861) : 289 (Rutilia). Holotype 9, InponEs1A (Moluccas, Batjan, A. R.
Wallace) : BMNH, London.

uzita Walker, 1849 : 860 (Dexia). Holotype 2, AUSTRALIA : BMNH, London.

The locality was cited in the original description as ‘New Holland ?’ but the specimen
must without doubt have originated from Australia.

valentina Macquart, 1843 : 279 (122) (Amphibolia). Holotype or syntypes g, AUSTRALIA :
MH, Lille.

For further detail on original material in Macquart’s collection in Lille see Crosskey

(1971 : 263).

variegata Bigot, 1874 : 461 (Fovmosia). Lectotype 2, by designation of Crosskey (1971 : 299),
AUSTRALIA : BMNH, London.

velutina Bigot, 1874 : 463 (Formosia). Lectotype 2, by designation of Crosskey (1971 : 299),
AUSTRALIA (Tasmania) : BMNH, London.

versicolor Brauer & Bergenstamm, 1889 : 171 (103) (Chrvysopasta). Lectotype g, by fixation
of Malloch (19286 : 616), AUSTRALIA (Western Australia, Swan River) : NM, Vienna.

The type-material of this species consists of a g and a 2 specimen with the same type-
data; Malloch (1928 : 616) referred to the male as ‘type’ and the female as ‘allotype’, and
cited the type-data, and his action is here accepted as providing a valid lectotype fixation.

victoriae Malloch, 1936 : 13 (Senostoma). Holotype g, AUSTRALIA (Victoria, Gisborne,
1g.1ii.1922, G. Lyell) : SPHTM, Sydney.

vidua Guérin-Méneville, 1843 : 273 (Rutilia). Syntypes 1g, 3 2, AUSTRALIA : lost.

violacea Macquart, 1851 : 198 (225) (Chetogaster). Holotype g, AUSTRALIA (‘céte orientale’,
probably New South Wales) : MNHN, Paris.

viridescens Enderlein, 1936 : 424 (Pancala, as variety of gemmata Enderlein, 1936). Lecto-
type 3, by present designation (p. 119), PAPUA NEw GUINEA (New Britain, Kinigunang,
C. Ribbe) : MNHU, Berlin.

viridescens Enderlein, 1936 : 403 (Chrysorutilia, as variety of caesia Enderlein, 1936). Holo-
type 9, AUSTRALIA (Queensland, H. Peters) : MNHU, Berlin.

viridicingens Enderlein, 1936 : 421 (Hega). Holotype 3, INpDoneEs1A (Moluccas, Batjan) :
MNHU, Berlin.

viridinigra Macquart, 1846 : 307 (179) (Rutilia). Lectotype 9, by designation of Crosskey
(1971 : 290), AUSTRALIA (Tasmania) : MNHN, Paris.

viridis Malloch, 1936 : 19 (Chaetogaster). Holotype g, AUSTRALIA (New South Wales, Com-
boyne, Chisholm) : SPHTM, Sydney.

Malloch described this species from the male, as stated by him (Malloch, 1936: 19,
line 1) before the key but not in the description; the holotype is J, not 2 as stated by Para-
monov (1968 : 367).

viriditestacea Macquart, 1851 : 190 (217) (Rutilia). Lectotype g, by designation of Crosskey
(1971 : 290), AUSTRALIA (Tasmania) : MNHN, Paris.
viridithovax Bigot, 1874 : 457 (Formosia). Nomen nudum (no later validation).
Bigot (oc. cit) attributed the name viridithorax to Macquart with the citation ‘Rutilia

140 R. W. CROSSKEY

id., Macq., Dipt. exot.’, but the name was not published by Macquart and remains a
nomen nudum attributable to Bigot.

viridiventris Crosskey, 1972 : (present paper) (Formosia (Formosia)). Holotype 3, SoLoMoN
IsLaNnpbs (Guadalcanal, Tapenanje, 10-30.ix.1953, J. D. Bradley) : BMNH, London.

vittata Macquart, 1855 : 126 (106) (Rutilia). Holotype 2, AusTRALIA (South Australia, Ade-
laide) : BMNH, London.

vivipara Fabricius, 1805: 309 (ITachina). Neotyped, by present designation (p.120), AUSTRALIA
(New South Wales, Barrington Tops, 9.iv.1949, S. J. Paramonov) : ANIC, Canberra.

volucelloides Walker, 1861b : 289 (Rutilia). Lectotype g, by present designation (p. 120),
InponesiA (Moluccas, Batjan, A. R. Wallace) : BMNH, London.

walkeri Paramonov, 1968 : 400 (Prodiaphania). Holotype 2, AUSTRALIA (Western Australia) :
AMNH, New York.

wilsoni Paramonov, 1950 : 524 (Amphibolia). Holotype g, AUSTRALIA (Victoria, Warburton,
14.xl1.1930, F. E. Wilson) : NMV, Melbourne (ex Wilson coll.).

[wilsont Paramonov, 1954 : 281 (Chaetogaster). Not Rutiliini as herein defined.]

zabirna Walker, 1849 : 863 (Dexia). Lectotype g, by present designation (p. 118), AUSTRALIA
(Western Australia, Perth, G. Clifton) : BMNH, London.

ACKNOWLEDGEMENTS

It is with much pleasure that I acknowledge the unfailing helpfulness of colleagues
in other museums and institutions who have generously provided me with types
and other material on loan. For such assistance I warmly thank the following:
Mr D. M. Ackland (formerly at University Museum, Oxford); Dr Donald Colless
(C.S.I.R.O., Canberra); Dr P. J. van Helsdingen (Rijksmuseum van Natuurlijke
Historie, Leiden); Dr A. Kaltenbach (Naturhistorisches Museum, Vienna) ; Professor
D. J. Lee (School of Public Health & Tropical Medicine, Sydney) ; Monsieur L. Matile
(Muséum National d’Histoire Naturelle, Paris); Dr D. K. McAlpine (Australian
Museum, Sydney); Dr Per Inge Persson (Naturhistoriska Riksmuseum, Stockholm) ;
Dr Curtis Sabrosky (U.S.D.A./U.S.N.M., Washington, D.C.); Dr H. Schumann
(Museum fiir Naturkunde der Humboldt-Universitat, Berlin); Dr J. Verbeke
(Institut Royal des Sciences Naturelles de Belgique, Brussels); Dr P. Wygodzinsky
(American Museum of Natural History, New York).

In addition I am very grateful to Monsieur Matile and to Dr L. Tsacas for their
generous help during a visit to the Muséum National d’Histoire Naturelle, Paris,
in September 1969, and to Mrs M. E. Crosskey for her invaluable help in making
copious notes on Macquart’s types during this visit. I thank Dr R. Defretin,
Conservateur of the Musée d’Histoire Naturelle in Lille, for information on specimens
in Macquart’s own collection at Lille.

REFERENCES

Austen, E. E. 1907. The synonymy and generic position of certain species of Muscidae
(sens. lat.) in the collection of the British Museum, described by the late Francis Walker.
Ann. Mag. nat. Hist. (7) 19 : 326-347.

Baer, K. E. von. 1827. Beitrage zur Kenntnis der niedern Thiere. Nova Acta physico-med.
[Bonn] 13 (2) : 523-762.

Baranoy, N. 1936. Weitere Beitrage zur Kenntnis der parasitaren Raupenfliegen (Tachi-
nidae = Larvaevoridae) von den Salomonen und Neubritannien. Ann, Mag. nat. Hist.
(10) 17 : 97-113.

7

REVISIONARY CLASSIFICATION OF RUTILIINI 141

Bezz1, M. 1928. Diptera Brachycera and Athericera of the Fiji Islands based on material in
the British Museum (Natural History). British Museum (Natural History), London, 220 pp.

Bicot, J. M. F. 1874. Diptéres nouveaux ou peu connus. 3° partie, IV. Genres Rutilia
et Formosia. Annls Soc. ent. Fr. (5) 4 : 451-467.

1880. Dipteres nouveaux ou peuconnus. 12 partie, XVIII. Genres Plagioceva (Macq.),
Formosia (Guérin) et Rutilia (Rob.-Desv.). Annls Soc. ent. Fr. (5) 10 : 85-89.

BRAvER, F. 1899. Beitrage zur Kenntnis der Muscaria schizometopa. Sber. Akad. Wiss.
Wien 108 : 495-529.

BRAUER, F. & BERGENSTAMM, J. E. von. 1889. Die Zweifliigler des kaiserlichen Museums
zu Wien. IV. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive
Anthomyidae). Denkschr. Akad. Wiss., Wien 56 (1) : 69-180.

& 1891. Die Zweifliigler des kaiserlichen Museums zu Wien. V. Vorarbeiten
zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Pars II.
Denkschr. Akad. Wiss., Wien 58 : 305-446.

— & 1893. Die Zweifliigler des Kaiserlichen Museums zu Wien. VI. Vorarbeiten
zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Pars III.
Denkschr. Akad. Wiss., Wien 60 : 89-240.

Cotitess, D. H. & McAtpine, D. K. 1970. Diptera (Flies), pp. 656-740 in The Insects of
Australia, xiii +- 1029 pp., Melbourne University Press.

CrosskKEY, R. W. 1965. A systematic revision of the Ameniinae (Diptera: Calliphoridae).
Bull. Br. Mus. nat. Hist. (Ent.) 16 : 33-140.

1967. An index-catalogue of the genus-group names of Oriental and Australasian
Tachinidae (Diptera) and their type-species. Bull. By. Mus. nat. Hist. (Ent.) 20 : 1-39.
— 1971. The type-material of Australasian, Oriental and Ethiopian Tachinidae (Diptera)

described by Macquart and Bigot. Bull. Br. Mus. nat. Hist. (Ent.) 25 : 251-305.

CurRAN, C.H. 1930. Four new Diptera from Australia. Am. Mus. Novit. No. 422 : 1-4.

Donovan, E. 1805. An epitome of the natural history of the insects of New Holland, New
Zealand, New Guinea, Otaheite, and other islands in the Indian, Southern, and Pacific Oceans :
etc. iv + 41 pls with descriptive text and index (unpaginated), London.

ENDERLEIN, G. 1922. Ein neues Tabanidensystem. Mitt. zool. Mus. Berl. 10 : 335-351.

1934. Dipterologica. I. Sber. Ges. naturf. Freunde Berl. 1934 : 416-429.

1936. Klassifikation der Rutiliinen. Verdff. dt. Kolon.-u. Ubersee-Mus. Bremen 1 : 397-
446.

EnceEL, E.O. 1925. Uber Rutiliidae sensu lat. (Dipt.). Zool. Jb. Abt. Syst. 50 : 339-376.

Ericuson, W. F. 1842. Beitrag zur Insecten-Fauna von Vandiemensland, mit besonderer

Beriicksichtigung der geographischen Verbreitung der Insecten. Avch. Naturgesch. 1842 (1) :
83-287.

| Fasricius, I. C. 1775. Systema entomologiae, sistens insectorum classes, ovdines, geneva

species, adiectis synonymis, locis, descriptionibus, observationibus. Flensburg & Leipzig,

832 pp.

1805. Systema antliatorum secundum ordines, geneva, species adiectis synonymis, locis,
observationibus, descriptionibus. Brunswick, 373 + 30 pp.

GERSTAECKER, A. 1860. Beschreibung einiger ausgezeichneten neuen Dipteren aus der
Familie Muscariae. Stettin. ent. Zig 1860 (4-6) : 163-202.

Gervais, P. 1838. Note sur la disposition systématique des Annélides chétopodes de la
famille des Nais. Bull. Acad. r. Sciences et Belles-lettves de Bruxelles 5 : 13-20.

Gray, G. 1832. In Cuvier, The Animal Kingdom arranged in conformity with its organization
15 (The class Insecta 2), London, 796 pp.

GUERIN-MENEVILLE, F. E. [1831]. In Duperrey, ed., Voyage autour du monde sur la corvette
de sa majesté La Coquille, pendant les années 1822, 1823, 1824-1825. Zoologie, Atlas,

Insectes. Paris, 21 plates.

1838. Crustacés, arachnides et insectes. In Duperrey, ed., Voyage autour du monde

sur la corvette de sa majesté La Coquille, pendant les années 1822, 1823, 1824-1825. Zool.

2 pt. 2, Div. 1. Paris, 319 pp.

142 R. W. CROSSKEY

GUERIN-MENEVILLE, F. E. 1843. Note monographique sur le genre de Muscides auquel
M. Robineau-Desvoidy a donné le nom de Rutilia, précédée de 1l’établissement d’un nouveau
genre voisin de celui-ci. Revue zool. 1843 : 262-274.

Macguart, J. 1843. Dipteres exotiques nouveaux ou peu connus. 2 (3) : Mém. Soc. Sci.
Agric. Lille 1843 : 162-460 (5-304).

1846. Diptéres exotiques nouveaux ou peu connus [1®] Supplément. MJém. Soc. Sci.

Agric. Lille 1844 : 133-364 (5-238). ;

1847. Diptéres exotiques nouveaux ou peu connus. 2¢ Supplément. Mém. Soc. Sci.

Agric. Lille 1846 : 21-120 (5-104).

1848. Diptéres exotiques nouveaux ou peu connus. Suite du 2™¢ Supplément [known

as 3rd Supplement]. Mém. Soc. Sci. Agric. Lille 1847 : 161-237 (1-77).

1851. Dipteéres exotiques nouveaux ou peu connus. Suite du 4® Supplément publié

dans les Mémoires de 1849. Mém. Soc. Sct. Agric. Lille 1850 : 134-294 (161-364).

1855. Diptéres exotiques nouveaux ou peu connus. 5° Supplément. Mém. Soc. Sci.
Agric. Lille (2) 1 (1854) : 25-156 (5-136).

Mattocu, J. R. 1927. Notes on Australian Diptera. No. xii. Pyroc. Linn. Soc. N.S.W.
52 : 336-353.

—— 1928a. Notes on Australian Diptera. No. xv. Proc. Linn. Soc. N.S.W. 53 : 319-335.

— 1928b. Notes on Australian Diptera. No. xvii. Proc. Linn. Soc. N.S.W. 53 : 598-617.

— 1928c. Notes on Australian Diptera. No. xviii. Pvoc. Linn. Soc. N.S.W. 53 : 651-662.

— 1929 Notes on Australian Diptera XX. Proc. Linn. Soc. N.S.W. 54 : 283-343.

—— 1930. Notes on Australian Diptera. XXIII. Proc. Linn. Soc. N.S.W. 55 : 92-135.

—— 1935. Insects Samoa 6 (Diptera) (9) : 329-366.

1936. Notes on Australian Diptera. XXXV. Proc. Linn. Soc. N.S.W. 61 : 10-26.

MEIJERE, J. C. H. DE. 1904. Note V. Zwei neue Dipteren aus dem Ostindischen Archipel.
Notes Leyden Mus. 24 (December 1903) : 177-178.

MEsniL, L. 1939. Essaisur les Tachinaires (Larvaevoridae). Monogrs Cent. natn. Rech. agron.
No. 7,66 + vpp. Paris.

OSTEN SACKEN, C. R. 1881. Enumeration of the Diptera of the Malay Archipelago collected
by Prof. Odoardo Beccari, Mr L. M. D’Albertis and others. Anmnali Mus. civ. Stor. nat.
Genova 16 : 393-492.

PaRAMONOV, S. J. 1950. Notes on Australian Diptera (I-V). Ann. Mag. nat. Hist. (12) 3:

515-534-

1954. Notes on Australian Diptera (XIII-XV). Ann. Mag. nat. Hist. (12) 7 : 275-297.
1968. A review of the tribe Rutiliini (Diptera: Tachinidae). I. Genera other than
Rutilia Robineau-Desvoidy and Formosia Guérin-Méneville. Aust. J. Zool. 16 : 349-404.

ROBINEAU-DESvoIDyY, J. B. 1830. Essai sur les Myodaires. Mém. prés. div. Sav. Acad.
Sct. Inst. Fr. 2 : 1-813.

RoNDANI, C. 1864. Dipterorum species et genera aliqua exotica revisa et annotata novis
nonullis descriptis. Avcho Zool. Anat. Fisiol. 3 (1) : 1-99.

SABROSKY, C. W. & CRossKEY, R. W. 1969. The type-material of Tachinidae (Diptera)
described by N. Baranov. Bull. By. Mus. nat. Hist. (Ent.) 24 : 27-63.

ScHINER, J. R. 1868. In Reise dey Osterreichischen fregatte Novara um die Evde in den Jahren
1857, 1858, 1859. Zool. 2 (1), Diptera, 388 pp. Vienna.

SNELLEN VAN VOLLENHOVEN, S.C. 1863. Beschrijving van eenige nieuwe soorten van Diptera.
Versl. Meded. K. Akad. wet. Amst. 15 : 8-18.

Swainson, W. 1840. In Swainson & Shuckard, On the history and natural arrangement of
imsects. 406 pp. London.

THomson, C. G. [1869]. Diptera. Species novas descripsit. In Kongliga svenska fregatien
Eugenies vesa omkving jorden. 2. Zool. 1 (Insecta) : 443-614. K. Svenska Vetenskaps-
Akademien, Stockholm ‘1868’.

Townsenp, C. H. T. 1915. Proposal of new muscoid genera for old species. Proc. biol. Soc.
Wash. 28 : 19-24.

REVISIONARY CLASSIFICATION OF RUTILIINI 143

TownsEND, C. H. T. 1916. Diagnoses of new genera of muscoid flies founded on old species.
Proc. U.S. natn. Mus. 49 (No. 2128) : 617-633.

1927a. Pyrodiaphania, new name for Diaphania Macquart (1843) preoccupied (Dipt.,

Muscoidea). Ent. News 38 : 159.

1927b. New Philippine Muscoidea. Philipp. J. Sct. 33 : 279-290.

1932. Notes on Old World Oestromuscoid types. Part II. Ann. Mag. nat. Hist.

(10) 9 : 33-57.

— 1936. Manual of Myiology. Part III. 255 pp. Itaquaquecetuba, Sao Paulo.

1938. Manual of Myiology. Part VII. 434 pp. Itaquaquecetuba, Sao Paulo.

— 1942. Manual of Myiology. Part XII. 365 pp. Itaquaquecetuba, Sao Paulo.

WALKER, F. 1849. List of the specimens of dipterous insects in the collection of the British

Museum 4 : 689-1172.

1859. Catalogue of the dipterous insects collected in the Aru Islands by Mr A. R. Wallace,
with descriptions of new species. J. Proc. Linn. Soc. Lond. 3 : 77-131.

—— 1861a. Catalogue of the dipterous insects collected at Dorey, New Guinea, by Mr A. R.
Wallace, with descriptions of new species. J. Proc. Linn. Soc. Lond. 5 : 229-254.

— 1861b Catalogue of the dipterous insects collected in Batchian, Kaisaa and Makian,
and at Tidon in Celebes, by Mr A. R. Wallace, with descriptions of new species. J. Proc.
Linn. Soc. Lond. 5 : 270-303.

— 1862. Catalogue of the dipterous insects collected at Gilolo, Ternate and Ceram, by
Mr A. R. Wallace, with descriptions of new species. J. Proc. Linn. Soc. Lond. 6 : 4-23.

1864. Catalogue of the dipterous insects collected in Waigiou, Mysol, and north Ceram

by Mr A. R. Wallace, with descriptions of new species. J. Proc. Linn. Soc. Lond. 7 : 202-

238.

: 1865. Descriptions of new species of the dipterous insects of New Guinea. J. Proc.

Linn. Soc. Lond. 8 : 102-130.

| WriepEemann,C.R.W. 1830. Aussereuropdische zweifliigelige Insekten. 2. 684pp. Hamm.

144 R. W. CROSSKEY

vertex

interfrontal parafrontal

area

eye

arista
antenna

parafacial

facial carina

epistome gena

occiput

postorbit

postbucca

genal dilation

2 3

Fics 1-3. Head of a Rutiliine fly. 1 & 2, laterodorsal and profile views showing the
terminology of main landmarks (vestiture omitted). 3, profile showing weak frontal
bristling and vibrissae of typical forms.

REVISIONARY CLASSIFICATION OF RUTILIINI 145

(/

6 Fi 8

Fics 4-8. Head profile, in outline only, of: 4, Rutilia, showing genal dilation (G.D.)
reaching to a level with anteriormost point ofeye. 5, Rutilodexia, showing feeble genal
dilation not reaching forwards nearly as far as anteriormost point ofeye. 6, Chryso-
pasta, facial outline only. 7, Prodiaphania, showing strong development of gular region
of head and short-plumose arista. 8, Chetogaster, facial outline only.

146 RK. W. CROSSKEY

humeral callus

humerals

posthumerals

prescutum
notopleurals

notopleuron presutural

seta

w—--—-=--—-—--O-

transverse —

suture supra-alars

scutum intra-alars

acrostichals

dorsocentrals

postalar callus postalars

a

supernumerary
prescutellars

preapical

scutell
scutellum telars

marginal
scutellars

10 1]

Fics 9-11. Thoracic dorsum of a Rutiliine fly. 9, terminology of sclerites (left side of
mid line) and chaetotaxy indicated by pores (right side of mid line). 10 & 11, posterior
view of scutellum showing level of insertions of marginal scutellar setae in Rutilia (10)
and Formosia (11). All figures schematic and relative bristle sizes indicated approxi-
mately by pore size shown (chaetotaxy very varied, forms may have more or fewer setae
in any series than indicated or may lack some series completely: see discussion of
taxonomic characters).

REVISIONARY CLASSIFICATION OF RUTILIINI 147

mentum

12 13 15

prosternal

prosternum membrane

edge of coxa

18

16 17

sternopleuron

19 20

/ Fics 12-20. Taxonomic characters used in the text. 12 & 13, two forms of proboscis
| found in Rutiliini: (12) mentum in profile with subparallel upper and lower edges, (13)
mentum in profile distinctly tapering apically. 14 & 15, two forms of buccal opening (oral
cavity) found in Rutiliini: (14) short broad buccal opening of most forms, (15) long
| narrow buccal opening of Prodiaphania and Foymodexia (both drawn from females:
buccal opening of males relatively slightly narrower in bothtypes). 16 & 17, vestiture
| of inner anterior surface of fore coxa in (16) Foymosia s. str. (almost whole surface haired)
and in (17) other Rutiliini (bare on inner part). 18, hairing of prosternal membrane
| found in many forms. 19 & 20, extent of hairing on pteropleuron in (19) subgenera
Chrysovutiia and Ameniamima, and (20) in other Rutiliini (hairing of sternopleuron
omitted).

148 R. W. CROSSKEY

postalar callus

scutum

suprasquamal
ridge

26

lower calypter

21

tegula

basicosta

24 25 27

Fics 21-27. Taxonomic characters used in the text. 21, postalar callus and surrounding
structures. 22-25, vestiture of postalar wall and suprasquamal ridge in: 22, forms with
both areas bare (e.g. Prodiaphania, Rutilodexia); 23, forms with sparse hairing only on
suprasquamal ridge (e.g. subg. Microrutilia); 24, forms with very long dense hair on
suprasquamal ridge (e.g. subg. Chrysorutilia, Amphibolia); 25, forms with dense hair
tuft on postalar wall (Formosia, Formodexia). 26, strongly explanate costal base (e.g.
Prodiaphania). 27, basal part of wing, showing long wiry setulae on tegula in most
forms (shaded area indicates the sub-basal dark mark found on the wing of most Ruti-
liini).

REVISIONARY CLASSIFICATION OF RUTILIINI 149

33 34 35

Fics 28-35. Abdominal characters. 28 & 29, the two main forms of abdomen in Rutiliini
with tergite numbering (pores indicate a typical chaetotaxy associated with each
abdominal shape). 30-35, fifth abdominal sternite of g in: 30 & 31, great majority of
forms; 32, subgenus Microrutilia; 33, subgenus Grapholostylum; 34 and 35, the two species
of subgenus Paramphibolia (34, stolida; 35, assimilis). Hairing omitted.

150 Rs W. CROSSKEY

epandrium
(T9)
37
cercus
surstylus
aedeagus
36 38
pregonite
epandrium
hypandrium

surstylus

distiphallus of aedeagus

epiphallus

postgonite

3?

Fices 36-39. Male genitalia of Rutiliini. 36, whole hypopygium in profile. 37 & 38, two
forms of aedeagal distiphallus found in Rutiliini: (37) great majority of forms in which
apical membranous part (A) is subequal in length to or shorter than basal sclerotized
part (B), (38) form in subgenus Grapholostylum in which membranous apical part whip-
like and nearly twice as long as sclerotized basal part. 39, aedeagus and associated struc-
tures viewed obliquely. All hairing omitted.

REVISIONARY CLASSIFICATION OF RUTILIINI 151

46 47

profile of Fovmosia fusca sp.n. 41-44, range of shape in profile of the surstylus in For-
mosia s. str. 45-47, epandrium, cerci and surstyli in profile (hairing omitted) of Cheto-

| Fics 40-47. Male genitalia in Formosia and Chetogaster. 40, complete hypopygium in
|
| gaster canberrae (45), C. oblonga (46) and C. violacea (47).

152

Figs 48-53.

Re Wo CROSSIGEY

Epandrium, cerci and surstyli of g genitalia in Formosia s. 1.
(left) and profile (right). 48, F. (F.) flavipennis. 409, F. (F.) viridiventris sp. n. 50, —
F. (Pseudoformosia) satuvatissima. 51, F. (P.) excelsa. 52, F. (Euamphibolia) speciosa. —
53, F. (E.) faceta. UWairing omitted.

Apical view

REVISIONARY CLASSIFICATION OF RUTILIINI 153

Fics 54-59. Epandrium, cerci and surstyli of f genitalia in Rutilias.1. Apical view (left)
and profile (right). 54, R. (R.) vivipava. 55, R. (R.) confusa. 56, R. (R.) dentata sp. n.
57, R. (R.) setosa. 58, R. (Ameniamima) argentifera. 59, R. (Neorutilia) simplex. Hair-
ing omitted.

154 R. W. CROSSKEY

64 65

Fics 60-65. Epandrium, cerci and surstyli of § genitalia in Rutilia s.1. Apical view
(left) and profile (right). 60, R. (Microrutilia) media. 61, R.(M.) nigripes 62,R (M.)
minor. 63, R. (M.) hirticeps. 64, R. (Grapholostylum) dorsomaculata. 65, R. (Dono-
vanius) agalmiodes. WHairing omitted, except for tuft on surstylus of minor.

REVISIONARY CLASSIFICATION OF RUTILIINI 155

70 71

Fics 66-71. Epandrium, cerci and surstyli of ¢ genitalia in subgenus Donovanius. Apical
view (left) and profile (right). 66, R. (D.) pellucens. 67, R. (D.) analoga. 68, R. (D.)
sabrata. 69, R. (D.) inusta. 70, R. (D.) vegalis. 71, R. (D.) lepida. Wairing omitted.

156 R. W. CROSSKEY

Fics 72-77. Epandrium, cerci and surstyli of ¢ genitalia in subgenus Chrysorutilia.
Apical view (left) and profile (right). 72, R. (C.) splendida. 73, R (C.) decova. 74,
R.(C.) crypticasp.n. 75, R.(C.) imperialis. 76, R. (C.) imperialoides sp.n. 77, RB. (C.)
idesa. UHairing omitted.

REVISIONARY CLASSIFICATION OF RUTILIINI 157

80 81

82 83 84

| Fics 78-84. Epandrium, cerci and surstyli of ¢ genitalia in subgenus Chrysorutilia.

| Apical view (left) and profile (right), and profile shape of surstylus only for some species.

| 78, R. (C.) rubriceps. 79, R. (C) caeruleata. 80, R. (C.) panthea. 81, R. (C.) formosa.
82, R. (C.) transversa, profile of surstylus. 83, R. (C.) caesia, profile of surstylus. 84,
R. (C.) chersipho, profile of surstylus. Hairing omitted.

158 R. W. CROSSKEY

Fics 85-90. Epandrium, cerci and surstyli of ¢ genitalia in Amphibolia and Rutilodexia
Apical view (left) and profile (right). 85, A. (Pavamphibolia) assimilis 86, A. (P.)
stolida. 87, A. (Amphibolia) ignovata. 88, A. (A.) valentina. 89, A. (A.) campbell.
90, Rutilodexia ? papua. Hairing omitted.

REVISIONARY CLASSIFICATION OF RUTILIINI 159

93

94

Fics 91-94. Epandrium, cerci and surstyli of ¢ genitalia in the genus Chrysopasta.
Apical view (left) and profile (right). 91, specimen from Newdegate, W.A. 92, speci-
men from 6 miles north of Watheroo, W.A. 93, holotype of elegans Macquart, locality

unknown. 94, lectotype of zabiyna Walker, Perth, W.A. (See text for discussion of
variability.) Hairing omitted.

£60 R. W. CROSSKEY

99 100

Fics 95-100. Epandrium, cerci and surstyli of J genitalia in Prodiaphania. Apical view
(left) and profile (right). 95, P: testacea. 96, P. furcata. 97, P. minuta. 98, P. avida.
99, P. fullerae. 100, P. genitalis. Main hairing only shown. oe

REVISIONARY CLASSIFICATION OF RUTILIINI 161

101
102

103 ~ 104 105

Fics 101-105. Epandrium, cerci and surstyli of g genitalia in Prodiaphania. to1, P.
georget, apical view and profile. 102, P. victoriae, apical view and profile. 103, P.
. funebris, semi-profile (drawn from old slide mount of 3 genitalia of paratype specimen
in which hypopygium not mounted exactly in profile). 104, P. deserta, profile. 105,
P. furcata, surstylus only in profile.

162 R. W. CROSSKEY

106 107

108 109

Fics 106-109. Epandrium, cerci and surstyli of ¢ genitalia in Prodiaphania and Rutiha.
Apical view (left) and profile (right). 106, P. biaymata. 107, P. cygnus. 108,
P. regina. 109, Rutilia (Microrutilia) nigriceps.

163

REVISIONARY CLASSIFICATION OF RUTILIINI

‘osuel [eqII}
ay} JO JU9}xe SpreMysea JsoyZANy oy} syuoseider1 voures ‘payddiys uorynqiystp peyoedsns
‘YORI PIOS UOINI}sIP UMOUY “TUTPNY oqsz oyz Jo uoINqrystp [eorydeis0an “AVI

= fears saatuy

a ik a Lh a BO OT T_T _
Ae: :

aN

{
x
St

/

/ J

yo

jane pias iff

= fff faY

164 R. W. CROSSKEY

INDEX TO GENUS-GROUP NAMES

The following index contains all generic and subgeneric names cited in the text, and the main
entry for each genus-group segregrate is shown in bold type.

Acucera, 18, 22

Agalmia, 18, 19, 22, 43, 83, 84, 129

Amenia, 51, 53, 83, 85

Ameniamima, 13, 14, 45, 47, 50, 51-54, 56,
71, 80, 85, 128

Amphibolia s.l., 18, 22, 24, 34, 39, 93-102,
103, 104, 108, 129

Amphibolia s. str., 95-100, ror, 129

Billaea, 21, 114
Bothrostira, 40, 41, 128

Chaetogaster, 112, 113, 130

Chaetogastrina, 94, 100, IOI, I13, 130

Chetogaster, mn, 133) 05, 1c; 19) 25, 22,23,
39, 94, Tor, 110, 112-116, 130

Chromocharis, 10, 25, 26, 33, 34, 127

Chrysopasta, 9, II, 15, 18, 22, 24, 39, 96,
tor, 102-107, 108, 130

Chrysorutilia, 9, 10, 13, 19, 39, 43, 45, 46,
47, 49, 50, 52, 53, 54-69, 70, 71, 80, 89,
035 Liz, £23, 1245, 125, 226

Codium, 112, 113, 130

Deximorpha, 22

Diaphania, 22, 107, I10, 130

Doleschalla, 18

Donovanius, 15, 17, 39, 40, 43, 44, 45, 46,
47, 49, 56, 69-77, 79, 80, 81, 92, 93, 123,
125, 128

Echrysopasta, 102, 107, 130
Euamphibolia, 25, 26, 32, 33-38, 96, 127
Euchaetogyne, 21

Euchrysopasta, 102

Eucompsa, 22, 43, 86, 87, 129

Formodexia, 15, 22, 23, 24, 25, 38-40, 94,
108, 128

Honmosta silo, 45) 10, 12 ens, ss) ieee
25-38, 42, 43, 44, 50, 53, 54, 79, 80, 94,
96, 127

Formosia s. str., 13, 26-31, 32, 34, 35, 127

Formosiomima, 53, 96

Formotilia, 22, 55, 59, 128

Grapholostylum, 16, 19, 43, 45, 46, 47, 50,
53, 50, 70, 71, 80, 83-86, 88, 89, 129

Habrota, 19, 43, 55, 117, 128

iHlega, 10; 25,260), 34, 34, 35, 227
Idania, 43, 55, 56, 58, 128
Laccura, 25, 26, 3I, 127

Menevillea, 43, 69, 70, 128
Microrutilia, 9, 16, 22, 43, 44, 45, 46, 47, 50,
84, 86-92, 93, I10, 129

Nemoraea, 6
Neorutilia, 43, 45, 47, 48-51, 57, 71, 80, 128

Ola, 6, 22, 114
Oxyrutilia, 6

Pancala, 25, 26, 32, 127

Paramphibolia, 16, 94, 95, 96, 100-102, 103,
I14, 129

Philippoformosia, 43, 55, 56, 58, 128

Pogonagalmia, 43, 86, 87, 129

Prodiaphania, 13, 14, 15, 16, 18, 22, 23, 39,
94, 107-112, 130

Prosena, 18

Prosenostoma, 43, 86, 87, 88, 129

Psaronia, 43, 69, 70, 72, 128

Psaroniella, 43, 70, 77, 79, 129

Pseudoformosia, 25, 26, 31-33, 35, 127

Rhynchiodexia, 108

Roederia, 18, 22, 102, 103, 130

Rutilia s.l., 4, 6, 11, 12; 18,22, 24,925,530)
40, 41, 42-93, 94, 95, 101, 108, IIO, 122,
128

Rutilia s. str., 14, 43, 45, 47, 54, 79 71, 72,
77-83, 123, 124, 129

Rutilodexia, 14, 21, 22, 24, 25, 39, 40-42,
94, 128

Rutilosia, 22, 40, 41, 128

Ruya, 6, 22, 114

Senostoma, 18, 86, 88, 107, 108, 110, 130
Stiraulax, 43, 77, 79, 129

Stomina, 21

Tachina, 77

Winthemia, 10, 46

Zoramsceus, 43, 55, 128

INDEX 165

INDEX TO SPECIES-GROUP NAMES

accedens, 130, 131

aditha, 73, 129, 131

agalmiodes, 72, 74, 128, 131, 154 (fig.)

albocincta, 96, 97, 99, 129, 131, 134

albopicta, 83, 84, 85, 129, 131

albovirida, 53, 85, 86, 129, 131

analoga, 72, 74, 128, 131, 155 (fig.)

angustecarinata, 131

angustigena, 62, 117, 128, 131

angustipennis, 40, 41, 128, 132

argentifera (Chetogaster), 115, 116, 130, 132

argentifera (Rutilia), 51, 52, 53, 54, 128, 132,
153 (fig.)

arida, 110, 112, 130. 132, 160 (fig.)

assimilis, 93, 94, 100, IOI, I14, 130, 132,
149 (fig.), 158 (fig.)

atribasis, 33, 34, 35, 36, 119, 127, 132

atrox, 55, 56, 58, 128, 132

atrox-group, 57, 58, 59, 128

auriceps, I15, 116, 130, 132

australasia, 62, 122, 128, 132

barcha, 73, 129, 132

biarmata, I10, 111, 130, 132, 162 (fig.)
bisetosa, 69, 70, 71, 72, 74, 119, 128, 132
blattina, 29, 31, 127, 132

bracteata, 29, 31, 127, 132

brevitarsis, 110, III, 130, 132
brunneipennis, 72, 75, 76, 128, 132

caeruleata, 61, 64, 65, 117, 128, 132, 157 (fig.)

caesia, 61, 63, 117, 128, 132, 157 (fig.)

callipygos, 26, 29, 31, 127, 132

campbelli, 94, 96, 97, 129, 132, 158 (fig.)

canberrae, 115, 116, 130, 132, 151 (fig.)

castanifrons, 72, 129, 132

castanipes, 70, 72, 77, 81, 129, 133

chersipho, 55, 61, 64, 122, 123, 128, 133,
157 (fig.)

cingulata, 32, 42, 44, 51, 52, 54, 128, 133

claripennis, I10, I1I, 130, 133

commoni (Amphibolia), 97, 129, 133

commoni (Prodiaphania), 110, III, 130, 133

complicita, 33, 35, 36, 127, 133

confluens, 62, 118, 128, 133

confusa, 42, 44, 54, 80, 81, 129, 133, 153 (fig.)

corona, 61, 63, 64, 128, 133

cryptica, 57, 60, 61, 62, 64, 65, 67, 128, 133,
156 (fig.)

cupreiventris, 90, 91, 129, 133

cygnus, 107, II0, 112, 130, 133, 162 (fig.)

decora, 60, 61, 65, 66, 67, 84, 122, 123, 124,
128, 133, 156 (fig.)

dentata, 81, 129, 133, 153 (fig.)

deserta, I10, 112, 130, 133, 161 (fig.)

desvoidyi, 72, 78, 80, 81, 119, 129, 133

diversa, 114, 133

dorsomaculata, 53, 83, 85, 86, 129, 133,
154 (fig.)

dubia, 133

dubitata, 72, 128, 133

durvillei, 80, 81, 123, 124, 129, 133

echinomides, 110, III, 130, 133

echinomyidea, 110

elegans, 96, 102, 103, 105, 106, 107, 130, 133,
159 (fig.)

engeli, 35, 36, 118, 127, 133

€0S, 29, 30, 118, 127, 133

erichsonii, 55, 61, 123, 128, 133

erronea, 130, 134

ethoda, 72, 75, 128, 134

evanescens, 62, 128, 134

excelsa, 32, 33, 127, 134, 152 (fig.)

faceta, 35, 37, 127, 134, 152 (fig.)

fervens, 29, 31, 127, 134

flavipennis, 29, 30, 127, 134, 152 (fig.)

flavipes, 86, 87, 88, 89, 90, I2I, 129, 134

formosa, 54, 55, 56, 59, 60, 61, 63, 122, 123,
124, 125, 128, 134, 157 (fig.)

formosa-group, 57, 58, 59, 128

formosina, 62, 128, 134

frontosa, 54, 128, 134

fulgida, 72, 73, 125, 129, 134

fullerae, 110, 112, 130, 134, 160 (fig.)

fulvipes (Enderlein), 29, 127, 134

fulvipes (Guérin-Méneville), 33, 34, 35, 122,
128, 134

fulviventris, 88, 90, 91, I21, 129, 134

funebris, 110, 111, 112, 130, 134, 161 (fig.)

furcata, I09, I10, III, 130, 134, 160 (fig.),
161 (fig.)

fusca, 35, 36, 37, 38, 127, 134, 151 (fig.)

fuscisquama, 85, 129, 134

fuscotestacea, 73, 129, 135

gemmata, 29, 31, 118, 127, 135

genitalis, 110, 111, 130, 135, 160 (fig.)
georgei, 109, I10, III, 130, 135, 161 (fig.)
glorificans, 29, 30, 127, 135

goerlingiana, 61, 63, 117, 128, 135
gratiosa, I15, 130, 135

166 INDEX

grisea, 130, 135

heinrichiana, 29, 30, 127, 135

heinrothi, 29, 31, 127, 135

hirticeps, 86, 87, 88, 89, 90, 129, 135, 154 (fig.)
humeralis, 130, 135

idesa, 61, 62, 128, 135, 156 (fig.)

ignobilis, 39, 40, 128, 135

ignorata, 96, 97, 99, 129, 135, 158 (fig.)

imitator, 73, 129, 135

imperialis (Amenia), 53

imperialis (Rutilia), 4, 60, 61, 64, 68, 69, 122,
123, 125, 128, 135, 137, 156 (fig.)

imperialoides, 62, 64, 67, 68, 69, 128, 135,
156 (fig.)

incomparabilis, 130, 135

inornata, 71, 72, 73, 80, 81, I19, 129, 135

inusta, 70, 72, 75, 76, I2I, 129, 135, 155 (fig.)

leonina (Amenia), 53

lepida, 73, 74, 122, 123, 125, 126, 129, 135,
155 (fig.)

leucosticta, 85, 129, 135

lineata, 61, 118, 128, 136

liris, 89, 90, 91, 129, 136

lucigena, 33, 120, 127, 136

luzona, 55, 56, 58, 59, 128, 136

luzona-group, 49, 55, 57, 58, 59, 128, 136

media, 90, 91, 92, 93, I10, 129, 136, 154 (fig.)

micans, 85, 86, 129, 136

micropalpis, 14, 15, 42, 47, 92, 93, 129, 136

minor, 22, 86, 87, 89, 90, 9I, 129, 136,
154 (fig)

minuta, I10, I12, 130, 136, 160 (fig.)

mirabilis, 4, 25, 26, 27, 29, 31, 127, 136, 137

mirabilis-group, 27, 29, 30, 127

moneta, 31, 32, 33, 127, 136

nana, 62, 128, 136

nigra, 73, 129, 136

nigribasis, 72, 118, 128, 136

nigriceps, 87, 90, 91, 129, 136, 162 (fig.)
nigrihirta, 71, 73, 75, 129, 136

nigripes, 90, 91, I19, 129, 136, 154 (fig.)
nigrithorax, 114, 136

nitens, 62, 123, 128, 136

oblonga, 112, I13, I1I5, I16, 130, 136,
151 (fig.)

obscuripennis, 33, 127, 136

onoba, 73, 129, 136

pallens, 90, 129, 136

panthea, 62, 63, 128, 137, 157 (fig.)

papua, 41, 42, 128, 137, 158 (fig.)

papuana, 95, 97, 98-100, 129, 137

paratestacea, 110, 130, 137

pauper, 32, 33, 119, 127, 137

pectoralis, 29, 127, 137

pellucens, 69, 70, 73, 74, 75, 122, 123, 126,
128, 137, 155 (fig.)

pellucida, 115, 116, 130, 137

plumicornis, 29, 127, 137

potina, 73, 129, 137

pretiosa, 34, 35, 120, 127, 137

prisca, 40, 41, 42, 128, 137

pubicollis, 61, 120, 128, 137

quadripunctata, 32, 53, 54, 128, 137

ralumensis, 41, 118, 128, 137

regalis, 4, 69, 70, 73, 74, 75, 122, 123, 126,
129, 137, 155 (fig.)

regina, I10, 112, 130, 137, 162 (fig.)

retusa, 4, 70, 73, 74, 125, 129, 137

rubriceps, 62, 63, 64, 123, 128, 137, 157 (fig.)

rufibarbis, 61, 118, 128, 137

ruficornis (Bigot), 61, 128, 137

ruficornis (Macquart), 89, 90, 91, I09, IIo,
129, 138

sabrata, 50, 70, 71, 72, 73, 74, 81, 129, 138,
155 (fig.)

sapphirina, 35, 127, 138

saturatissima, 31, 33, 120, 127, 138, 152 (fig.)

savaiiensis, 71, 73, 76, 129, 138

scutellata, 92, 93, 129, 138

semifulva, 61, 128, 138

serena, 62, 123, 128, 138

setosa, 70, 77, 81, 82, 122, 123, T20;eE2o)
138, 153 (fig.)

sexpunctata (Amenia), 83

simplex, 44, 48, 49, 50, 51, 128, 138, 153 (fig.)

smaragdifera, 35, 127, 138

smaragdina, 35, 37, 128, 138

solomonicola, 29, 31, 127, 138

soror, 130, 138

speciosa, 33, 34, 35, 36, 53, 94, 120, 128, 138,
152 (fig.)

spinipectus, 73, 120, 129, 138

spinolae, 73, 129, 138

splendida (Donovan), 4, 57, 59, 60, 62, 64,
65, 66, 67, 84, 122, 123, 124, 125, 128, 132,
138, 139, 156 (fig.)

splendida (Townsend), 55, 56, 58, 59, 128,
138

INDEX 167

stolida, 94, 96, 100, IOI, 102, 113, I14, 130,
138, 149 (fig.), 158 (fig.)

subtustomentosa, 84, 85, 86, 129, 138

subvittata, 61, 128, 138

testacea, 7, 107, 108, 109, 110, III, 130, 138,
160 (fig.)

townsendi, 55, 59, 128, 139

transfuga, 71, 73, 76, 129, 139

transversa, 62, 65, 128, 139, 157 (fig.)

trixoides, 39, 40, 128, 139

uzita, 61, 128, 139

valentina, 7, 93, 94, 95, 96, 97, 129, 139,
158 (fig.)

variegata (Rutilia), 85, 129, 139

variegata (Senostoma), 88, 108

velutina, 85, 129, 139

versicolor, 102, 103, 105, 106, 107, 130, 139

victoriae, I10, III, 112, 130, 139, 161 (fig.)

vidua, 97, 122, 129, 139

Dr R. W. CrosskeEy, D.Sc., A.R.C.S., F.I.Biol.

Department of Entomology

British Museum (Natura History)
CROMWELL Roap

Lonpon, SW7 5BD

violacea, I12, 113, I15, 116, 130, 139,
151 (fig.)

viridescens (Chrysorutilia), 61, 128, 139

viridescens (Pancala), 29, 119, 127, 139

viridicingens, 33, 35, 127, 139

viridinigra, 70, 73, 129, 139

viridis, 115, 116, 130, 139

viriditestacea, 73, 129, 139

viridithorax, 130, 139

viridiventris, 26, 28, 30, 127, 140, 152 (fig.)

viridiventris-group, 27, 28, 127

vittata, 110, III, 130, 140

vivipara, 42, 72, 77, 78, 80, 81, 119, 122, 123,
124, 126, 127, 129, 140, 153 (fig.)

volucelloides, 38, 39, 40, 120, 128, 140

walkeri, 109, I10, III, 130, 140
wilsoni (Amphibolia), 96, 97, 98, 129, 140
wilsoni (Chaetogaster), 114, 140

zabirna, 102, 105, 106, 107, 118, 130, 140,
159 (fig.)

“o Gay Aes |. Get Bae bees pT 4 448 oe ced 105 OE
OP 424. wer SE. C5 hwy q
i (> jdt 7? timege Oy seacestink> mo. Cam glia
er @ 44.24 ee ee ee © ot A, :

et diet See eer ee oe ws
WES gto) fy eee. Stkatt a te 2M ey

) Pei rei he be iu he ¥ 9
1 0? See QF ern eotege . oes ee Sd
~— —— iy | bY eee eas ‘es 235-0, Die

( ieee if eatin (hain : oh £2 geea hen ee

. ae ss pe“) (Arr ; > > Caeptst ae

. i 7
. . “a. » é Py & age os: £5 A) _
i. * =
1 pls . a - wee ott te a

oe)
a)

1-2: se Aol wT? php seg ees ast hee

A RECLASSIFICATION {sccm
OF THE SUBFAMILY AGRYPNINAE »,,, .<°
(COLEOPTERA: ELATERIDAE) |

C. M. F. voN HAYEK

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
_ ENTOMOLOGY Supplement 20
LONDON : 1973

~~

me RECTUASSIFICATION OF ‘THE SUBFAMILY. .. )
maORYPNINAE (COLEOPTERA: ELATERIDABy,, «

BY

CHRISTINE MARIA FELICITAS VON HAYEK

Pp. 1-309; 17 Text-figures

BULLETIN. OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 20
LONDON ? 1073

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted im 1949, %s
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at regular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be
completed within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Supplement 20 of the Entomological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. nat. Hist. (Ent.). Suppl.

© Trustees of the British Museum (Natural History), 1973

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 18 October, 1973 Price £12.30

A RECLASSIFICATION OF THE SUBFAMILY
mGRYPNINAE (COLEOPTERA: ELATERIDAE)

By CHRISTINE MARIA FELICITAS von HAYEK

CONTENTS
Page
SYNOPSIS ; : ; : E : : ; : : 3 3
INTRODUCTION ; - : “ 4
THE CLASSIC CHARACTERS USED FOR THE SEPARATION OF THE GENERA
THE CHARACTERS USED FOR THE SEPARATION OF THE GENERA IN THE
PRESENT WORK : 5
NOTES ON THE KEY TO GENERA AND SUBGENERA . 5
KEY TO GENERA AND SUBGENERA ‘ : : ; : ; , 6
THE ASSIGNMENT OF SPECIES TO GENERA : ; 8
TYPE-MATERIAL AND THE DESIGNATION OF LECTOTYPES : : : 9
NOTES ON THE FORMAT OF THE ANNOTATED CATALOGUE. ; : : 10
ABBREVIATIONS AND SYMBOLS USED IN THE CATALOGUE ; : , 10
NOTE ON THE LINE AS A UNIT OF MEASUREMENT. : 3 : : II
SPECIES REMOVED FROM THE SUBFAMILY AGRYPNINAE . ; ‘ P II
SPECIES INCERTAE SEDIS . : , 12
THE PROPOSED CLASSIFICATION OF THE "AGRYPNINAE, WITH AN ANNOTATED
CATALOGUE OF THE KNOWN SPECIES. ; : : ; 12
CHECKLIST OF THE GENERA INCLUDED IN THE AGRYPNINAE ; ‘ : 268
ACKNOWLEDGMENTS : : : : , : 269
SOME NOTES ON THE COLLECTIONS CONSULTED. : ; ; ; 270
REFERENCES . : f ‘ ‘ ; F 4 : : 2 281
INDEx . : : : : ‘ : : : : : : 297
SYNOPSIS

The characters until now used to separate the genera of Agrypninae are reviewed and
evaluated and the use of new and previously neglected characters is proposed. The type-
material of the type-species of every genus except Adelocera was located and examined. The
genera are redefined on the basis of the new characters. A key to the genera is provided.

The diagnostic features of the type-species of Adelocera are deduced from the lectotype of
another species, shown, from a study of the literature and other evidence, to be conspecific
with it.

Eight hundred and ninety-five of the nominal species included in the Agrypninae by previous
workers are evaluated and each is assigned to an appropriate genus. Four hundred and
seventy-one new combinations are proposed. Thirty-five generic and 23 specific names are
newly placed in synonymy. One generic and 11 specific names are extracted from synonymy.
The location of the type-material (holotypes and syntype-series) of 115 species is recorded and
406 lectotypes are newly designated.

4 C;. Me Bs von EAWEK

INTRODUCTION

THE Elateridae considered in this work are those comprising the Agrypninae of the
Schenkling Catalogue (1925), with the exception of Anaspasis (see p. 5), and
those subsequently described as belonging or related to the genera in this subfamily.
The genus Octocryptus is included as a matter of convenience as the species bear
a superficial but close resemblance to certain Agrypninae.

The category subfamily is adopted for the group principally because it is the
one used in the most recent practical key to the Elateridae of the world (Fleutiaux,
19g41a : 36). Other workers have treated the same group of species as a tribe
(Agrypnini Schwarz, 1906, Adelocerini Arnett, 1969), subtribe (Agrypnina Hyslop,
IgI7 : 236) or intermediate group (Agrypnitae Fleutiaux, 194Ic).

Recent years have seen the establishment of a number of new genera and
subgenera belonging to the Agrypninae. A study of the literature has shown that
in almost every case the diagnosis of the old established and new genera and
subgenera are little more than brief summaries of the most conspicuous features
of the type-species. In very few cases only are there any indications of the range
of variation which may be found within the genus or any effort made to define its
limits. Attempts to identify material has shown that many species cannot be
accommodated satisfactorily in any genus as at present defined, while others can
be placed equally well in two or more genera.

This unsatisfactory state of affairs suggests that the characters until now used
for the separation of the genera display a greater degree of variation than has
been hitherto suspected. The reason why this has only now become apparent is
that both past and present workers have based their studies on restricted faunas
and shown little or no interest in the relationship of the species considered in their
works with those found in other regions. In addition, workers have continued
to use the same classic characteristics for the separation of the genera as their
predecessors over a hundred years ago, and almost no attempt has been made
to evaluate these characters at the generic level or to discover new ones.

THE CLASSIC CHARACTERS USED
FOR THE SEPARATION OF THE GENERA

The characters until now used for the separation of the genera have been the
length and depth of the tarsal grooves, the appearance of the surface and lateral
margins of the prothorax and elytra, the form of the mesosternal groove, the length
of the antennae and of the femora of the middle legs and the structure of the fourth
tarsal segment.

Examination of a great deal of material in the British Museum (Natural History),
the Muséum National d’Histoire Naturelle, Paris, the Institut Royal des Sciences
Naturelles de Belgique, Brussels, and other collections has shown that the degree
to which these characteristics are developed varies from one species to another
and sometimes from one individual to another and also that they occur in such
varied combinations that they are of no value at the generic level. However the
present investigation and a study of the literature has led to the discovery of new

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 5

and previously neglected characteristics which are of considerable importance at
the generic level.

THE CHARACTERS USED FOR THE
SEPARATION OF THE GENERA IN THE PRESENT WORK

The most important of the characters used in the present work is the structure
of the middle coxal cavity. Thomson (1864 : 59, 1868 : 91), Schiodte (1865 : 497) and
more recently Nakane & Kishii (1956 : 202, 203) were aware that this character was of
some importance, but as Kontkanen (1964 : 48) comments, it has been completely
forgotten by taxonomists. The other main characteristics are the size and shape
of the second and third antennal segments, the presence or absence of tibial spurs,
the position of the propleural grooves for the reception of the tarsi in relation
to the antennal groove and of the distal end of the metasternal groove for the
reception of the tarsi in relation to the margin of the metasternum.

NOTES ON THE KEY TO GENERA AND SUBGENERA

With the aid of the following key there should be no difficulty in assigning to the
appropriate genus, as defined in this paper, all species listed in the subfamily
Agrypninae in the Schenkling (1925) catalogue, with the exception of those
belonging to the genus Anaspasis (see below), and any Agrypnine species described
after the publication of the catalogue.

With one exception (Elater ovalis Germar — see below) the characters used in the
key have been observed in the holotype or lectotype specimen of the type-species
of each genus. In only one case (that of Elater ovalis Germar, the type-
species of Adelocera) did the search for the type-material or indeed any determined
material prove completely unsuccessful. In this case the generic characters are
drawn from the lectotype of another species, Lacon occidentalis Candéze, shown
later in the present work (p. 21) to be congeneric with, and very similar in
appearance to, ovalis Germar.

Anaspasis Candéze (1882 : 4) was excluded from the Agrypnitae [sic] by Fleutiaux
(1941c : 48) because of the strong resemblance it bears to Protelater Sharp (1877).
Golbach (1953 : 278) is in agreement with Fleutiaux. Examination of the type-
species of the two genera, Protelater elongatus Sharp (1877 : 482) and Anaspasis
fasciolata Candéze (1882: 5), has shown that Fleutiaux was fully justified in his
action. Both species differ from all Agrypnine genera in the shape of the frons
and the absence of an antennal groove occupying a position corresponding to the
whole or part of the length of the prosternopleural suture. In addition they lack
setae at the base of the claws and possess tibial spurs. Crowson (1961 : 160)
discusses the apparent affinities of Protelatery but the true relationships of this and
certain other genera, including Anaspasis, have yet to be discovered.

Fleutiaux (1907) : 170) included the monobasic genus Aniws Candéze (1889) in
the ‘Agrypnides’ but omits it without comment from his key to the Agrypnitae
(Fleutiaux, 1941c). The type-material of the type-species, gracillimus Candéze,
was received on loan shortly before this paper was submitted for publication.
The inclusion of the species in the Agrypninae has been confirmed but further

6 C. M. BY von HAYEK

studies of additional material (when this becomes available) are required before its
relationships with other species can be elucidated.

KEY TO GENERA AND SUBGENERA

I Propleurae with longitudinal grooves near the lateral margin for the accommodation
of the antennae. Posterior portion of prosternopleural suture grooved for the
reception of the anterior tarsi : : : OCTOCRYPTUS Candéze (p. 266)

— Propleurae without longitudinal grooves near the lateral margin for the accom-
modation of the antennae. Prosternopleural suture never modified for the
reception of the tarsi

ii 8

Fics 1-8 Figs 1-3. Mesocoxal cavity and adjacent area. 1. Lanelatey sp. 2. Agvypnus
sp. 3. Adeloceva sp. Figs 4-6. Meso- and metasternal area, left middle leg removed.
4. Opatelus sp. 5. Agrypnus sp. 6. Adelocera sp. Figs 7-8. Left side of prosternum and
propleuron, front leg removed. 7. Agrypnussp. 8.Adelocerasp. All figures are diagramatic
and are not drawn to the same scale. Mesosternal groove and mesocoxal cavities stippled,
tarsal grooves cross-hatched. EMz,mesepimeron. ES2,mesepisternum. S2, mesosternum.

$3, metasternum.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 7

NN

Mesepisternum forming part of margin of middle coxal cavity (Text-fig. 1). Tibial
spurs present (Text-fig. 9). Vestiture setose, scales entirely absent
LANELATER Arnett (p. 240)
— Mesepisternum not forming part of margin of middle coxal cavity (Text-figs 2 & 3).

Tibial spurs absent (Text-fig. 10). Vestiture generally scaly : ; : : 3
3 Mesepimeron not forming part of margin of mesocoxal cavity (Text-fig. 2) . : 4
— Mesepimeron forming part of margin of mesocoxal cavity (Text-fig. 3) . : 8
4 Tarsal segments 1-4 expanded laterally (Text-fig. 17) . TRIERS Candéze (p. 239)
— Tarsal segments 1-4 not expanded laterally (Text-figs 15 & 16) . : 5

5 Small species, less than 5mm long. Prothorax constricted immediately pened tie

anterior angles. Lateral carina not attaining the anterior margin of the prothorax
RISMETHUS Fileutiaux (p. 235)

— Larger species or, if less than 5 mm long, the prothorax not constricted behind the

anterior angles and lateral carina attaining the anterior margin of the prothorax 6
6 Scutellum with longitudinal carina 2 2 . : : : - : 7
— Scutellum simple, without carina . : - AGRYPNUS Eschscholtz (p. 113)
7 Propleurae without tarsal grooves . : MERISTHUS Candéze s. str. (p. 231)

— Propleurae with tarsal grooves
MERISTHUS subgenus SULCIMERUS Fileutiaux (p. 231)
8 Claws without basal setae (Text-fig. 12) . : . DANOSOMA Thomson (p. 87)
— Claws with basal setae (Text-fig. 13) . é : ‘ ; : : 9
9 Second and third antennal segments subequal (Text-fig. 13). If the third is
triangular and larger than the second, then propleural tarsal grooves are present
and run parallel to the antennal grooves 5 : : ‘ : ‘ ; 10

14

15 16 Ve

Fics 9-17. Fig. 9. Lanelatey sp., apex of tibia, pubescence partially removed. Fig. to.
Agrypnus sp., apex of tibia. Fig. 11. Agrypnus sp. claw. Fig. 12. Danosoma sp., claw.
Figs 13-14. Antennal segments one to four. 13. Agrypnussp. 14. Laconsp. Figs 15-17.
Tarsal segments. 15. Agrypnus sp. 16. Agrypnus sp. 17. Trievs vamitarsus Candéze.

8 C.. MLS Ee VON VELA

— Second and third antennal segments not subequal (Text-fig. 14). The third
segment as long as the fourth but sometimes of a different shape : 12
10 Propleurae and metasternum with well defined grooves for the reception of the
tarsi. Metasternal grooves directed posteriorly; the distal end of these grooves,
if produced, would cut the posterior margin of the metasternum (Text-fig. 4)
OPATELUS Candéze (p. 91)
Propleurae and metasternum with or without grooves or depressions for the
reception of the tarsi. Metasternal tarsal grooves or depressions, if present,
directed laterally; the distal end, if produced, would cut the lateral margin of
the metasternum at a point at or within the anterior three-quarters of its length
(Text-fig. 6) : : : : Il
11 Lateral margins of pessooaa with a band of Glosely packed seales
SCAPHODERUS Candéze (p. 51)
— Lateral margins of pronotum without a band of closely packed scales
ADELOCERA Latreille (p. 13)

12 Tarsal segments without ventral lobes (Text-fig. 15) . ‘ : : 13
— One or more tarsal segments with ventral lobes (Text-fig. 16) : 3 : F 15
13 Lateral margins of prothorax not carinate . ; : EIDOLUS Candéze (p. 190)
— Lateral margins of prothorax carinate : ‘ 14

14 Posterior prosternal process directed upwards immediately belied the anterior coxae
CANDANIUS nom. n. (olim Anius Candéze) (p. 85)
— Posterior gota process not directed upwards immediately behind the anterior
coxae : : . LACON Castelnau (p. 52)
15 Abdomen with Sree ia the eonoed mE the pare tarsi
ACROCRYPTUS Candéze (p. 92)

— Abdomen without grooves for the reception of the hind tarsi : 16
16 African species. Antennal groove not extending beyond the ee half a the
prosternopleural suture : 17

— American and Indonesian species. Proehoracie Pp ieaeal aoe eee faced
the anterior half of the prosternopleural suture DILOBITARSUS Latreille (p. 93)
17 Prothorax cylindrical. Head with a pair of tubercles directed anteriorly. Frons
not margined anteriorly ; ; : . ELASMOSOMUS Candéze (p. 101)
— Prothorax not cylindrical. Head without tubercles. Frons margined anteriorly
HEMICLEUS Candéze (p. 108)

THE ASSIGNMENT OF SPECIES TO GENERA

The major part of this work is concerned with the assignment of each Agrypnine
species known to me to the appropriate genus. It also provides a record of the
present location of the type-material of the majority of these species.

With a few exceptions, generic assignments have been made only for those species
known to me from examinations of the holotype, paratype, lectotype or syntype
material, or specimens whose identification labels or provenance leads me to believe
that they are correctly identified.

The generic assignment of species by means of diagnostic characters unknown
to earlier workers, renders it impossible to make generic assignments for the
majority of species known to me only from the description. In the few cases in
which the describer has recorded characters of secondary importance, such as
the position of the propleural tarsal grooves, a generic assignment has been made.
Generic assignments based on descriptions must be regarded as tentative until they
have been confirmed by examination of the type-material.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 9

EVE E-MATERETAL AND THE DESIGNATION OF LECTOTYPES

Until comparatively recently it was not generally accepted practice amongst
workers on the Elateridae, when they described a new species, to designate a single
specimen as the holotype. Many early writers did not even record the number
of specimens they had before them at the time of the description or to whom they
belonged or where they were preserved. The majority did no more than place
a determination label on one or more specimens. Sometimes they included the
word ‘type’ on the label, but no worker appears to have been consistent in his
actions.

Many subsequent workers and museum curators have accepted these specimens
with the describer’s determination label (either with or without the word ‘type’)
as holotypes and very often affixed distinctive type-labels (referred to as ‘curatorial
labels’ in the present work) to them.

While the acceptance of these ‘types by tradition’ may appear to be perfectly
satisfactory, in the majority of cases it can also create considerable difficulties.
These arise mainly because in the past the word ‘type’ did not have the same
meaning as it does at the present time. It was often used by workers to indicate
that, in their opinion, a particular specimen was a typical example of a species.
As a result several specimens, sometimes belonging to different species or even
different genera and preserved in different collections, have been labelled and
treated as holotypes.

In order to overcome these difficulties Article 73 of the International Code of
Zoological Nomenclature has been strictly observed. In the absence of a definite
statement by the author that the description of a new species is based on a single
specimen or that a particular specimen is the type (or some similar unambiguous
expression) it has been assumed that the description is based on a syntype series
and that a lectotype designation is required.

In this work lectotypes have been designated for those species for which it has
been possible to locate and assemble all the known extant specimens of the original
series or at least the greater part of it. Wherever possible the specimen marked
‘type’ by the describer, or that traditionally regarded as the type, whether so
labelled or not, has been designated as the lectotype.

Where a reviser has stated that a particular specimen is the type, when it is
clear that the original description is based on more than one specimen (e.g. Van
Zwaluwenburg, 1959 : 354, Lacon variolus Candéze [=Agrvypnus, see p. 228] described
from an unrecorded, but large, number of specimens) this statement has been
accepted as a valid lectotype designation unless there is sufficient evidence that
the description was based on another specimen or the specimen is unsuitable for
some other reason (e.g. badly damaged or the genitalia lost).

It has been assumed, unless the author states that the material on which the
description is based is the property of some other individual or institution, or is
part of a collection made by travellers or exploring parties and subsequently
acquired by a collector or institution, that the type-material was in the describer’s
own collection. Syntype material found in other collections is assumed to have
been acquired by the new owner as a gift or by exchange or purchase. Short

Io Ci Me EBs von HAYEK

notes on the history of each collection consulted in the course of this work will be
found on pages 270-281.

NOTES ON THE FORMAT OF THE ANNOTATED CATALOGUE

Except in the case of Adelocera, which is treated in greater detail (see p. 22), the
following information is given for each species: the original combination with the
author, date and reference and, where there has been a change, the most recently
published combination. In those cases in which a previously published combination
is restored, the first use of that combination is recorded.

Labels on holotypes, paratypes, lectotypes and paralectotypes are quoted
verbatim throughout, but in addition the country name (when known for certain)
is given first in its current form, in capitals, whether or not it occurs on the labels.
All lectotypes, paralectotypes and syntypes and also, where necessary, holotypes
and paratypes have been appropriately labelled. Labels on other specimens are
quoted verbatim in sufficient detail for them to be identified where this is necessary;
in other cases the data are listed in a standardized form. The handwriting on
labels has been identified wherever possible, and the name of the writer given in
square brackets. Labels without this information are in a handwriting unknown
to me or printed.

Geographical place-names appearing in The Times Index-Gazetteer of the World,
1965, are recorded without comment. For place-names not appearing in The Times
Index-Gazetteer the alternative present-day name or the latitude and longitude are
given in square brackets. A list of gazetteers consulted in the course of this work
will be found at the end of the References.

ABBREVIATIONS AND SYMBOLS USED IN THE CATALOGUE

; A semicolon is used to show the extent of each label where this is quoted
verbatim.

/ An oblique line indicates the extent of data on upper and lower side of one label.

[ } Square brackets indicate additional information concerning labels, such as the
identification of handwriting, the meaning of numbers, letters or signs on, or
the colours of labels, alternative spellings of place-names, etc.

? A question mark before the name of a collection indicates that the available
evidence indicates that the material in question is believed to be in that
collection but that its presence there has not been confirmed.

The names of institutions, museums and collections containing material are
abbreviated as follows:

AM, Sydney Australian Museum, N.S.W., Australia.

ANS, Philadelphia Academy of Natural Sciences of Philadelphia, Pennsylvania,
U.S.A.

BMNH British Museum (Natural History), London, England.

BPBM, Honolulu Bernice P. Bishop Museum, Hawaii.

CAS, San Francisco California Academy of Sciences, U.S.A.

CIE, London Commonwealth Institute of Entomology, c/o BMNH.

CM, Pittsburgh Carnegie Museum, Pennsylvania, U.S.A.

CMSN, Milan Civico Museo di Storia Naturale, Italy.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE Il

DEI, Eberswalde

FRI, Dehra Dun
HSPA, Honolulu
IMZU, Turin
IRSNB, Brussels
IZPAN, Warsaw
MCSN, Genoa
MCZ, Harvard
MM, Brno

MN, Santiago
MNHN, Paris
MNHU, Berlin
NM, Basle

NM, Nairobi
NM, Vienna
NMV, Melbourne
NR, Stockholm
RNH, Leiden
SAM, Adelaide
TM, Budapest
TU, Turku

UM, Oxford
USNM, Washington

UZM, Copenhagen
ZI, Leningrad
ZIMLU, Halle-Wittenburg

ZM, Amsterdam
ZMU, Helenski
ZMU, Moscow
ZSBS, Munich
ZUM, Uppsala

Deutsches Entomologisches Institut, East Germany (formerly
at Berlin—Dahlem)

Forest Research Institute, India.

Hawaiian Sugar Planter’s Association, Hawaii.

Museo ed Istituto di Zoologia dell’Universita, Italy.

Institut Royal des Sciences Naturelles de Belgique, Belgium.
Instytut Zoologiczny, Polska Akademia Nauk, Poland.

Museo Civico di Storia Naturale, Italy.

Museum of Comparative Zoology, Cambridge, Mass., U.S.A.
Moravské Museum, Biologiky Ustav, Czechoslovakia.

Museo Nacional de Historia Natural, Chile.

Museum National d’Histoire Naturelle, France.

Museum fiir Naturkunde der Humboldt-Universitat, Germany.
Naturhistorisches Museum, Switzerland.

National Museum, Kenya. Formerly Coryndon Museum.
Naturhistorisches Museum, Austria.

National Museum of Victoria, Australia.

Naturhistoriska Riksmuseum, Sweden.

Rijksmuseum van Natuurlijke Historie, Netherlands.

South Australian Museum, Australia.

Természettudomanyi Museum, Hungary.

Department of Zoology, Turku University, Finland.

Hope Department, University Museum, England.

Smithsonian Institution, United States National Museum,
Washington, D.C., U.S.A.

Universitetets Zoologiske Museum, Denmark.

Zoological Institute of the Academy of Sciences, U.S.S.R.
Zoologisches Institut, Martin Luther Universitat, Halle-Witten-
burg, East Germany.

Zodlogisch Museum, Netherlands.

Zoological Museum of the University, Finland.

Zoological Museum of the University of Moscow, U.S.S.R.
Zoologische Sammlung des Bayerischen Staates, West Germany.
Zoologisk Universitets Museum, Sweden.

NOTE ON THE LINE AS A UNIT OF MEASUREMENT

Some nineteenth-century workers used the line as a unit of measurement in
their descriptions. A six inch ivory ruler produced by Janson of Russell Street,
London, shows that the English, French and German lines differ in length. In
the present work the following metric equivalents are used:

I English line = 2-117 mm
I German line = 2-IgI mm

1 French line = 2:250 mm

SPECIES REMOVED FROM THE SUBFAMILY AGRYPNINAE

Propsephus tropicus (Hope) comb. n.
Agrypnus tropicus, Hope, 18430 : 365.
LECTOTYPE (present designation), LrBERIA: 2 tropicus Hope, Palmas [Hope];
Type, Hope, Ann. Nat. Hist 11. 1843 p. 365. coll. Hope Oxon [UM, Oxford

I2 C2 MS EX von AY Ek

curatorial label]; Type coll. 15464, Agrypnus tropicus, Hope, Hope Dept [UM,
Oxford curatorial label] (UM, Oxford).

Paralectotype. 2 Palmas [Hope] and UM Oxford curatorial labels as above
except that this specimen bears the number 15462 (UM, Oxford).

The published locality is ‘West Africa, circa Palmas.’ [Cape Palmas.] Candéze
(1857 : 30) states that the species may be a synonym of A. puber Candéze (1857 : 30)
or a related species but that Hope’s description is so short that it is impossible to
recognize the species from the description. Examination of the type-material has
shown that Hope was mistaken in his generic attribution.

P. tropicus bears a very strong resemblance to P. beninensis (Candéze) but further
study of both species is necessary before the synonymy can be confirmed.

SPECIES INCERTAE SEDIS

The type-material of the following species cannot be found and is assumed to
be lost or destroyed. The descriptions are too poor to allow even a tentative
generic attribution to be made.

Lacon adanensis Jagemann
Lacon adanensis Jagemann, 1944 : 333.

The description is based on an unrecorded number of specimens from Adana
[TURKEY]. The Jagemann collection is now in the MM, Brno, but neither the
type-material nor any determined specimens can be found there. It is not in
Jagemann’s possession (Dr Stehlik in litt.).

Jagemann compares adanensis with ‘L. cyprii Baudi’. Baudi did not describe
a species of this name. Jagemann is presumably referring to L. pygmaeus Baudi,
1871, described from Cyprus (see p. 45).

Lacon anathesinus Candéze
Lacon anathesinus Candéze, 1897 : Io.

The description is based on an unrecorded number of specimens from CHINA.
The type-material cannot be found in the BMNH or IRSNB, in Brussels. No
determined specimens of this species which, according to Candéze, resembles
Anathesis laconoides Candéze, have been located.

THE PROPOSED CLASSIFICATION OF THE AGRYPNINAE,
WITH AN ANNOTATED CATALOGUE OF THE KNOWN SPECIES

Subfamily AGRYPNINAE Candéze

Agrypnides Candéze, 1857 : 17. Type-genus: Agrypnus Eschscholtz, 1829.
Agrypnites Jacquelin du Val, 1859 : 125.

Agrypnini Kiesenwetter, 1863 : 230.

Agrypnina Thomson, 1864 : 59.

Agrypnidae Fleutiaux, 1891 : 387.

Adelocerini Buysson, 1893 : 18. Type-genus: Adeloceva Latreille, 1829.
Adelocerinae Fleutiaux, 1926 : 92.

Agrypnitae Fleutiaux, 1941c : 42.

Adelocerina Blackwelder, 1944 : 280

Agrypninae Fleutiaux, 1947 : 241.

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 13

Certain recent workers (Arnett, 1952; Arnett, Mignot & Smith, 1969) base the
family-group name (subfamily, tribe), which they apply to the Elateridae considered
in the present work, on Adelocera. They are presumably basing their action on
Buysson (1893), who believed that a family-group name should be based on the
oldest included genus within the category under consideration. However Article 36
of the International Code of Zoological Nomenclature makes it quite clear that the
family-group name must be based on Agrypnus, on which Candéze (1857), who
was the first worker to propose a name for a supra-generic group, based the name
of the tribe Agrypnides. Candéze’s comment at the bottom of page 14 suggests
that Lacordaire was responsible for the naming of the tribe. Though this may
be true (Candéze, 1857 : vi), Candéze’s work bears the date Mai, 1857 whereas
Lacordaire’s did not appear until June of the same year (see p. 291).

Diacnosis. Underside of the prothorax with grooves for the reception of part, or the

whole, of the antennae. The grooves occupy the position and at least one third of the length
of the prosternopleural sutures. Meso- and metasternum not connate between the mesocoxae.

DISTRIBUTION. The subfamily is represented in all regions. The largest number
of genera and species is found in the Ethiopian and Oriental regions. Only a
single species, Agrypnus variabilis (Candéze), occurs in New Zealand. Sharp
(1877 : 479) comments that he suspects that this species has been introduced
[from Australia] by marine traffic; it appears now to be fairly well established
(see p. 227).

Though the characteristics listed above distinguish the species considered in this
work from all other Elateridae, I believe that the subfamily includes species which
are only very distantly related. Further studies and the evaluation of the impor-
tance at family-group level of such characteristics as the setae at the base of the
claws, the tibial spurs and the structure of the margin of the mesocoxal cavity may
eventually provide a basis for the subdivision of the family on more natural lines.

ADELOCERA Latreille

Adeloceva Latreille, 1829 : 451. Type-species: Elatey ovalis Germar, by subsequent designation
(Hyslop, 1921 : 632).

[Lacon sensu Germar, 1840 and subsequent authors nec Castelnau, 1836. Pars.]

Agraeus Candéze, 1857: 165. Type-species: Agraeus mannerheimi Candéze, by monotypy.
Syn. n.

_ Pericus Candéze, 1857 : 167. Type-species: Pericus nitidus Candéze, by monotypy. Syn. n.

Brachylacon Motschulsky, 1858 : 60. Type-species: Brachylacon microcephalus Motschulsky,
by monotypy. [Wrongly synonymized with Lacon Castelnau by Gemminger & Harold,
1864: 1491.] Syn.n.

Tvachylacon Motschulsky, 1858: 61. Type-species: Tvachylacon fulvicollis Motschulsky, by
subsequent designation (Hyslop, 1921 : 672). [Synonymized with Agvaeus Candéze by
Schwarz, 1906 : 27.]

| Cavicoxum Pic, 1928 : 21. Type-species: Cavicoxum monstrosum Pic, by monotypy. [Synony-

mized with Agraeus Candéze by Fleutiaux, 1931 : 79.]

Prolacon Fleutiaux, 1934d : 179. Type-species: Pyolacon alluaudi Fleutiaux by monotypy.
Syn. n.

Aganolacon Ohira, 1967 : 55 (as a subgenus of Brachylacon). Type-species: Aganolacon shirozui
Ohira, by monotypy. Syn. n.

14 Cc. M. F. von HAYEK

GENERIC DIAGNOsIS. In the absence of the type-material or any other specimens
of Elater ovalis Germar, the lectotype of Lacon occidentalis Candéze is used as the
basis for determining the fundamental diagnostic characteristics of the genus
Adelocera Latreille (see pp. 16-21).

Each claw bearing a group of setae near the base (Text-fig. 11). Tibial spurs absent
(Text-fig. 10). Mesepisternum does not form part of margin of mesocoxal cavity. Mesepimeron
forms part of margin of mesocoxal cavity (Text-fig. 3). Second and third antennal segments
subequal, each smaller than the fourth and following segments (Text-fig. 13). Antennal
groove not extending beyond the anterior half of the prosternopleural suture, deep enough
to accommodate the rolled antennae. Vestiture scale-like at least in part. Propleural tarsal
grooves, if present, run parallel to the antennal grooves (Text-fig. 8). Metasternal tarsal
grooves, if present, directed laterally so that the distal ends, if produced, would cut the laterai
margin at a point at or within the anterior three-quarters of its length (Text-fig. 6). Scutellum
variable in shape but never carinate. Tarsi with or without ventral lobes (Text-figs 15, 16).

RANGE OF VARIATION FOUND WITHIN THE GENUS. Species possessing this
combination of diagnostic characteristics display a wide range of variation in the
degree of development and modification of the anterior angles, lateral margin and
discal area of the prothorax, the appearance of the elytra, the depth of the
propleural and metasternal tarsal grooves, the structure of the fourth tarsal segment
and the distribution of scales and setae on the body. In some species these
characteristics are more strongly developed in one sex than in the other (see Sexual
Dimorphism, p. 21). Up to the present time these features have been used to
establish a number of genera and subgenera. Examination of material from all
parts of the world has demonstrated the existence of so many intermediate forms
that, in my opinion, there is no justification for the retention of the genera and
subgenera which are discussed below.

HIsTORY OF THE GENUS. The genus Adelocera was established by Latreille
(1829 : 457) for Elater ovalis Germar, Elater fuscus Fabricius ‘et quelques autres
[unspecified] des Indes Orientales raportés par M. de Labilliardiere’. In a post-
humous work Latreille (1834 :144) states that Adelocera ‘a pour types une espéce
de Java qui me parait étre l’Elater fuscus de Fabricius et un autre de Savanah,
trés raprochée de son E. marmoratus, si ce n’est pas lui.’ This statement is
unacceptable and invalid as the type-designation of the type-species of Adelocera.
Hyslop (1921 : 629) subsequently designated Elater ovalis Germar as the type-
species of A delocera.

Some years later Germar (1840 : 255) redefined Adelocera, presumably basing
his interpretation on Latreille, 1834, since he credits the genus to Latreille, and
includes marmoratus Fabricius and also lepidoptera Panzer and conspersa Gyllenhal,
varia Olivier and pennata Fabricius, fasciata Linnaeus and senilis Germar, but not
the species included in Adelocera by Latreille in 1829. Germar transferred E. ovalis
to Lacon. He does not mention Elater fuscus Fabricius at all, probably because
Eschscholtz (1829 : 32) had transferred it (erroneously, see p. 158) to Melanotus.
Thomson (1859 : 103) subsequently designated Elater fasciatus Linnaeus as the
type-species of Adelocera, which he credits to Latreille though there can be no
doubt that his concept of the genus is that of Germar.

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 15

Germar’s interpretation of Adelocera was generally accepted until Hyslop
(1921 : 621) drew attention to the fact that [as a result of his designation of species
originally included in the genera Adelocera Latreille, 1829 and Lacon Castelnau,
1836 to replace the type-species erroneously selected by Thomson (1859 : 103, see
above and p. 54)] ‘the two genera Adelocera and Lacon have been reversed in their
application’. Fleutiaux (19250) independently came to the same conclusion and
the following year he published a correction (Fleutiaux, 1926) to the Schenkling
catalogue to this effect. As Hyslop does not give any clear indication which
species are to be included in his interpretation of Adelocera and Lacon, the present
author has accepted Fleutiaux (1926) as the author and date of the new
combinations.

Unfortunately the majority of workers appear to have overlooked Hyslop’s
and Fleutiaux’s corrections, with the result that there are many apparently
conflicting generic attributions in the more recent literature. This, together with
the fact that the type-material of Elater ovalis cannot be found and that as a
result each worker has his own interpretation of the genus, is the main cause of
the confusion within the subfamily.

NOTES ON THE GENERA AND SUBGENERA PLACED IN SYNONOMY WITH Adelocera.
Agraeus was erected for a single species, mannerheimi Candéze. Candéze (1857 : 166)
remarks that the genus is based more on general appearance than ‘charactéres
réelment générique’. The present investigations have shown this to be true. The
characters used by Fleutiaux (1927 : 88 and 1935a@: 12) in his redefinition are all
of the variable type discussed on p. 14. The species included by these authors
display a considerable degree of sexual dimorphism.

Pericus. Candéze’s comment on Agraeus also applies to this monobasic genus.
Thirty-five years later (1892c : 485) he remarked that discedens Candéze is a ‘forme
de transition’ between Pericus and his interpretation of Lacon [=Agrypnus of the
present work], and that, in this species the generic characters ‘tendent de se
dégénérer pour rapprocher des Lacon du group II, div. C.’ (Candéze, 1891c : 23,
obesus Candéze, nebulosus Candéze, etc.). The species which were included in
this genus display a moderate degree of sexual dimorphism.

Brachylacon. The type-species, microcephalus Motschulsky, differs from the
present interpretation (see p. 38) of ovalis Germar, the type of Adelocera, only in
the depth of the propleural and metasternal tarsal grooves and the structure of the
fourth tarsal segment.

Trachylacon was erected to accommodate species differing from Brachylacon in
the puncturation of the elytra. In species attributed to Tvachylacon the punctu-
ration is confused and not arranged in regular rows. However, in certain species,

| such as fulvicollis Motschulsky, the elytral puncturation appears confused but

closer inspection shows that there is a definite tendency towards a regular striate
. in the apical portion of the elytra. The characters used by Fleutiaux
(19352) to redefine the genus are all of the variable type discussed on p. 14.

Cavicoxum. Pic erected the family Cavicoxumidae for this monobasic genus.

16 CMa ES von HAW Bik

Fleutiaux (1931 : 74) recognized it as a member of the Elateridae and at that
time regarded it as congeneric with Agrvaeus. Ten years later (1941c) he re-erected
the genus using the length of the lateral carina of the prothorax, which attains the
anterior margin, as the characteristic separating it from his interpretation of
Agraeus. The length of the lateral carina is one of the characteristics displaying
too great a range of variation to be of value at the generic level. The type-species,
monstrosus Pic, is the female of constrictus Ritsema, a species displaying a very
marked degree of sexual dimorphism.

Prolacon was erected for a single species, alluaudi Fleutiaux, which the author
described as having antennal grooves open anteriorly and widely excavate
posteriorly and metathoracic episterna which are parallel-sided and very narrow.
Examination of the type-material (see p. 32 under fleutiauxt) has shown that the
antennal grooves do not differ from those found in other species assigned to A delocera;
the prosternopleural suture is deeply grooved anteriorly and closed posteriorly. It
would appear that Fleutiaux misinterpreted the paired carinae on, and the steeply
declivous sides of, the posterior portion of the prosternum as forming the posterior
part of the antennal groove. The metathoracic episterna are visible only as very
small triangular sclerites lying beside the anterior angles of the metasternum. It
seems probable that Fleutiaux was misled by the carina which runs parallel and
close to the inner margin of the epipleurae. The size of the metathoracic episterna
does not appear to be of importance at the generic level. Fleutiaux also regarded
the tubercles on the prothorax and elytra and the humeral carinae as diagnostic.
Tubercles are discussed above. Humeral carinae are known to occur in other
Adelocera species, including trifasciatus microcephalus Motschulsky.

Aganolacon was erected as a subgenus of Brachylacon for those species in which
the tarsal grooves of the propleura are not distinctly margined. This is one of the
very variable characteristics discussed on p. 14.

THE TYPE-MATERIAL OF Elater ovalis GERMAR. Germar described Elater ovalis
from Persia on two occasions. On the first (1824: 49) he used the name Elater
ovalis Knochii [sic] but on the second (1840: 261) he omitted the reference to
Knoch. On neither occasion did he record the number of specimens he had before
him or the collection to which they belonged.

Fleutiaux (194Ic : 46) records that the type of Elater ovalis Germar is in the
ZMHU, Berlin. Dr F. Hieke, curator of Coleoptera in that institution has searched
the collections on my behalf without success. There are no specimens which are
named as, or which could be, ovalis Germar in the ZMHU, Berlin collection, which
contains part of the Germar and also all the known extant part of the Knoch
collection (see p. 276). Fleutiaux was in the habit of recording the fact that he
had seen the type of a species in some other collection by placing a note to this
effect in his own collection. The fact that there is no note on ovalis in his collection
suggests that he did not see the type-material but based his statement on the
assumption that the ovalis material was preserved in Germar’s own collection.

Germar’s collection was broken up (see p. 275) and in addition to the part in the
ZMHU, Berlin, there are portions in the DEI, Eberswalde, the ZIMLU, Halle-

;

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 17

Wittenburg (formerly the University Zoological Museum, Halle) and the BMNH
(see below). Dr Diekmann of the DEI, Eberswalde and Dr Hiising of the ZIMLU,
Halle-Wittenburg both write that there are no specimens of ovalis Germar in their
collections.

Candéze’s (1857 : 132) statement that he received material of ovalis from the
Germar collection ‘en communication’ [? = loan] from Schaum suggests that at
the time the type-material of ovalis was in Schaum’s possession. However the
ovalis material does not appear to have been included in that part of the Schaum
collection which was acquired by the BMNH, as there are no specimens in the
collection and the species is not listed in the BMNH manuscript catalogue of the
Schaum collection. If Candéze (see below) did return the ovalis material to Schaum,
it must have formed part of that portion of the Schaum collection which has been
lost (see p. 280).

If, either by accident or design, Candéze retained the ovalis material he received
from Schaum, it should have passed to the BMNH or the IRSNB, Brussels (see
p. 271). Neither collection contains any specimens determined as, or which
could be, ovalis Germar. The species is not recorded in Waterhouse’s manuscript
catalogue of Candéze’s Elateridae in the Janson collection in the BMNH.

Since the most careful and repeated searches through the relevant collections
have failed completely to discover the type-material of Elater ovalis Germar there
is no alternative but to accept the fact that, almost certainly, the material has
been lost or destroyed.

At first sight the loss of the type-material of Elater ovalis Germar, together with
the fact that no determined specimens of the species can be found (see also p. 21),
appears to make it impossible to draw up a generic diagnosis of Adelocera. How-
ever I believe that the evidence produced by the study of the literature is sufficient
to justify the use of the lectotype of Lacon occidentalis Candéze for this purpose.

THE DIAGNOSTIC FEATURES OF Elater ovalis GERMAR. Germar’s (1824 : 49)
Elater ovalis is a dark-coloured beetle with reddish legs and antennae. Germar
describes it as follows [translated from the Latin by CMFH]. ‘The round head is
pitchy black in colour and deeply sunk into the prothorax which is strongly
emarginate anteriorly. The prothorax is the same colour as the head but the

_ lateral margins are paler; it is convex with the lateral margins arcuate and slightly

convergent anteriorly; the posterior margin is straight and the posterior angles
more or less right-angled. The elytra are convex, as broad as the prothorax and
twice as long, the apex rounded. The head is closely punctured, the prothorax
closely and sparsely punctured and the elytra punctate-striate with convex
interstriae. The prothoracic pubescence is grey and the elytra bear squamiform
setae which arise from the punctures. The underside is black and coarsely and
closely punctured. The serrate antennae are accommodated in grooves on the
| underside of the prothorax, which also bears deep, shiny, angled depressions for the
reception of the anterior legs’. Germar does not record the length but states that
it is ‘shorter, but not narrower than E£. ripario’. Germar does not mention
E. npario in his 1824 publication. His first reference to the species (as E. riparius
B

18 C. MM. FS von HAY EK

Fabricius) is in 1844 (p. 139) when he published what amounts to a revision of
Cryptohypnus.

As the comparison with FE. vipario provides a clue to the size of E. ovalis it is
necessary to discover whether at that time there was a generally accepted interpre-
tation of E. ripario. In 1824 the only published descriptions of E. riparius were
those of Fabricius (1792 : 232 and 1801 : 243, referring to 1792), Paykull (1800 : 41,
referring to Fabricius, 1792) and Gyllenhal (1808 : 402, referring to Fabricius,
1801). The E. riparius figured by Panzer (1796) with a reference to Fabricius
(1792) was redescribed as mvularis by Gyllenhal (1808: 403). Schenkling
(1925 : 202) lists both species under Hypnoidus. The specimens standing under
these names in the BMNH, IRSNB, Brussels and MNHN, Paris indicate that
there is a general agreement concerning the identity of these two species, which
range in size between 4-7 mm and 7:5 mm.

In his second, shorter description of ovalis, Germar (1840: 261) refers to his
work of 1824. He gives the length as a little over 2 lines (about 5-5 mm, see p. II)
and remarks that the deep femoral and tibial grooves of the prothorax have sharply
defined margins. Germar does not otherwise amplify his original description.
He does, however, include ovalis in Lacon (sensu Germar not Castelnau, 1836,
see p. 14) so that it is not unreasonable to assume that ovalis possesses the
characteristics of his interpretation of that genus. Germar (1840 : 260) redefined
Lacon, transferring all the originally included species (see p. 53) to other genera.
According to Germar the characteristics of the genus Lacon are as follows:—

Short serrate antennae in which the 2nd and 3rd segments are small and the 12th more or
less invisible.1 The median portion of the prosternum is strongly protruberant and separated
from the lateral portions by grooves which extend the whole length of the sternite. [By this
Germar presumably means from the anterior margin of the prosternum to a point immediately
in front of the anterior coxae. In his description of L. caliginosus Boisduval, Germar comments
that this species differs from the others in the genus in that these grooves extend only a little
over half way.] Transverse grooves for the accommodation of the anterior femora are present
at the base of the lateral portion of the underside of the prothorax [propleurae] and, in some
species, oblique grooves for the reception of the anterior tarsi.

No known species possess this combination of characters and also the grooves
for the accommodation of the anterior tarsi mentioned in the description of ovalis.
The species which bear the closest resemblance to the description of ovalis are those
belonging to the genus Opatelus Candéze, 1857. In these Brazilian and Bolivian
species the antennal groove ends close to the anterior coxae, the distance between
the end of the groove and the coxa being equal to the diameter of the coxae.
When examined under a hand lens the groove appears to attain the coxa. Opatelus
species have lobed tarsi, but this is not very obvious under a hand lens.

It is however unlikely that Germar had a South American species before him
when he described ovalis. As Germar’s first description of ovalis refers to Knoch,

1 The antennae are presumably 11-segmented. In his description of mustelinus, Germar states that
the 12th antennal segment is visible. In this species (see p. 188) the apical third of the eleventh segment
is marked off from the rest of the segment by a constriction. This may have led Germar to believe that

the antennae were 12-segmented. The same is true of Agrypuus notodenta Latreille (see p. 256), which
according to Germar (1840 : 251, 252) has 12-segmented antennae.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 19

it seems reasonable to assume that he based his description on material in the

Knoch collection or on specimens received from Knoch. At the beginning of

the nineteenth century South American material was still something of a rarity

and as Zincken (1818) makes no mention of any such material in his account of

Knoch’s collection it is probable that none was present. The belief that ovalis is

not a South American Opatelus species is supported by the fact that Candéze

(1857 : 132), who examined Germar’s ovalis material, did not recognize it as a

member of his own new genus.

In the absence of any known species or undetermined material agreeing with
Germar’s description of ovalis, the only course left open is to consider the possibility
that Germar misinterpreted certain structural characteristics of the species included
in his interpretation of Lacon and that as a result his generic diagnosis is inaccurate
and misleading. This is not beyond the bounds of possibility, as some of the
included species are small and the position taken up by the legs when the beetles
are killed tends to obscure the details of the structure of the underside. Germar’s
optical equipment also will have been very inadequate by present-day standards.
It was probably no more than a simple hand-lens. It is also possible that the
inflammation of the eyes which afflicted Germar during the last ten years of his
life (he died in July, 1853) was already causing his eyesight to deteriorate (Schaum,
1853 : 380).

In order to discover whether this is the case, Germar’s material of his Lacon
species was, as far as possible, located and examined. The details of the specimens
are recorded in the catalogue portion of this work. The results of this investigation
are presented in the following table.

From the table it is immediately obvious that all Germar’s Lacon species differ
from his generic diagnosis in that the antennal groove does not extend the entire
length of the prosternum, or in other words it does not attain the anterior coxae.
In this characteristic all the species resemble caliginosus, which Germar regarded

as an exception, remarking that the antennal grooves extend only a little way
beyond the middle of the length of the prosternum. In all twelve known species
the tibial spurs are not merely indistinct but entirely absent.

It is not unreasonable to assume that ovalis possesses the same characteristics as
the other twelve species included in the genus. Evidence that this assumption
is justified is found in the work of Candéze (1857 : 130), who appears to have been
the last man to see Germar’s material. Candéze places ovalis in his interpretation
of Lacon, which differs from that of Germar in that the prosternopleural suture
is closed posteriorly so that the antennal groove does not attain the anterior coxae.

In the same work Candéze (1857 : 130) described Lacon occidentalis from Senegal,
remarking that this species could at first sight be confused with ovalis Germar
but can be distinguished by the presence of white scales among the black ones
clothing the body. The lectotype of L. occidentalis Candéze (a synonym of A delocera
| parcus (Boheman), see p. 43) is in the BMNH.

Additional information concerning the structure and appearance of ovalis is
| provided by Baudi (1871 : 49), who refers to ovalis Germar in his description of
| Lacon pygmaeus from Cyprus, remarking that his species may be little more than

|

20 C. M. F. von HAYEK

Table to illustrate how the species included in Lacon Germar, 1840 (nec Castelnau,
1836) agree and differ from the generic diagnosis
Other

Generic characteristics listed by Germar _character-
istics

or

(a)

Antennal segments 2 and 3 small, subequal
Propleurae with transverse grooves for the

Prosternopleural antennal extends the entire
accommodation of the femora

length of the prosternum
Mesepimeron forms part of margin of

Propleural tarsal grooves present (p)
mesocoxal cavity

Antennae 11-segmented* (2)
Tarsi simple, without ventral lobes

Tibial spurs present* (3)

Species included by Germar

caliginosus Boisduval (page 136) v v 4) v vp x v No
crenatus Klug sensu Germar (page 145) Vv v x v vp x v No
crenicollis Ménétriés (page 145) v v x v va x v No
tvvovatus Klug (page 171) v v x v va x v No
murinus Linnaeus (page 186) v v x Vv va x v No
mustelinus Germar (page 188) v v x v va x v No
muticus Herbst (page 189) v v x v vp x v No
nodifer Klug (page 191) v v x v va x v No
parviceps Schoenherr (page 196) v v x v vp x x No
tervvenus Germar [=crenatus Klug] v v x v vp x v No
(page 145)
turbidus Germar (page 224) v v x v va x v No
vestitus Klug (page 228) . , yt v x v va x No
ovalis Germar (page 16)
Species believed to resemble ovalis
Germar
occidentalis Candeze (page 43) . mag, v x v vp x v Yes
pygmaeus Baudi (page 45) . yh v x v Vp x v Yes

Key to the symbols used in the table
v = specimen agrees with Germar’s generic diagnosis.
x = generic character listed by Germar absent.
® = differs from generic diagnosis but agrees with the description of the species.

* (2) In the key Germar (1840 : 251) states that the 12th antennal segment is not visible. In the
generic diagnosis (page 260) he states that the 12th segment is more or less hidden.

* (3) Germar’s ‘kaum sichtbaren Enddorn’ suggests that the tibial spur is present but indistinct.

;

|

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 21

a variety of ovalis Germar. The type of pygmaeus is in the IMZU, Turin (p. 45).
Apart from the colour it bears a close resemblance to occidentalis Candéze. Whether
Baudi based his interpretation of ovalis on the description or some material, now
lost, is unknown.

The diagnostic characteristics of occidentalis Candéze and pygmaeus Baudi are
included in the table. It can be seen that they possess the characteristic features
of Germar’s Lacon species and differ from Germar’s generic diagnosis in the same
way as the other species.

These investigations have shown that no known species possesses the combination
of characteristics attributed to Lacon ovalis by Germar. Although it is conceivable
that such a species may yet be found, the evidence obtained from the re-examination
of Germar’s Lacon material suggests that Germar misinterpreted the structure
of the antennal groove in all the species (with the exception of caliginosus) included
in his interpretation of Lacon, and that as a result his generic diagnosis is inaccurate
and misleading. The antennal groove does not occupy the entire length of the
prosternopleural suture as stated by Germar, but extends only a little beyond
the half-way mark. As this type of groove also occurs in occidentalis Candéze
I consider that Candéze’s comment that ovalis bears a close resemblance to
occidentalis can be accepted without reservation.

OTHER SPECIMENS OF Elater ovalis GERMAR. Up to the present time I have
been unable to discover any specimens determined as ovalis Germar in any of the
museums and institutions listed on pages Io and II.

Latreille (1829 : 251) and Castelnau (1840: 24) are the only workers, apart
from Candéze, Baudi (see p. 19) and cataloguers, to make any reference to ovalis
Germar. As they do not record any specimens and as neither of their collections
contain determined material of ovalis it seems probable that they based their
interpretation of the species on the description.

There are no published records of the capture of ovalis from the type-locality,
Persia (Iran) or from anywhere else. Enquiries of the principal museums at home
and abroad have failed to discover any undetermined material from Persia which
bears a strong resemblance to occidentalis. Examination of the descriptions of
unknown species from South-west Asia attributed to Lacon sensu auct. (nec Castel-
nau) or related genera has failed to disclose any occidentalis-like species. Lacon
adanensis Jagemann (1944 : 333), described from Adana (Turkey), is approximately
the right size but the description is too poor to be of any value and up to the present
time it has proved impossible to find the type-material (see p. 12).

SEXUAL DIMORPHISM. Pronounced sexual dimorphism is not common in the
Elateridae. It is however very strongly developed in certain Adelocera species,
a fact that has not been recognized before and which has led to the males and
females of certain species being described as distinct species and in some cases
being attributed to different genera.

The sexual dimorphism resembles that found in other groups of insects, including
the Coleoptera, in that individuals of one sex are more ornate and often larger
than those of the opposite sex. Adelocera species are unusual in that it is the

22 CC. M. Fo von” HAMWEK

female which displays a greater degree of development and modification of the
prothorax. The smooth patch, pore or brush of setae present on the terminal
and/or penultimate abdominal sternites of the females of many Agrypnus species
does not occur in Adelocera.

DIsTRIBUTION. Up to the present time one species, (minutus (Candéze)) has
been recorded from the Nearctic while the remainder occur in the Ethiopian and
Oriental regions. All the Australian species listed under Adelocera by Neboiss
(1961) following Van Zwaluwenburg (1959) have been found, on examination,
to belong to the genus Agvypnus.

BIOLOGY AND HABITS. Nothing is known of the history and habits of Adelocera
species. The adults have been found on plants and trees and on the ground.

SPECIES INCLUDED IN THE GENUS

The following 75 species are now assigned to the genus.

Because of the unusual type of sexual dimorphism displayed by certain species
the genus has been studied in greater detail than the other genera included in
this work.

Every reference to each species has been consulted. In every case in which a
worker has recorded a specimen or specimens, every effort has been made to locate
and examine the material. Where this has proved possible, the reference is included
in the specific synonymy of the appropriate species and the data on the material
and its present location recorded. A list of any previously unrecorded specimens
is also given.

Adelocera adspersus (Candéze)

Lacon adspersus Candéze, 1857 : 115.
Adelocera adspersus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Inp1A: 9, N. India, Bacon; Janson coll.
1903 : 130; adspersus Cdz. type [Cand.]; Lacon adspersus Cdze., Cand. Type
e coll. de Laferté [Janson] (BMNH).

Candéze records the locality as ‘Des Indes-orientales-boreales’ (see p. 271).
I have not found any specimens with this locality. The discrepancy between
the published locality and the label is almost certainly due to Janson (see p. 276).

Candéze (1890 : cxcix and 1892c : 484) recorded this species from Chota Nagpore,
(see p. 273). The Fleutiaux collection in the MNHN, Paris contains a female
which I believe to be part of Candéze’s series. It bears Fleutiaux’s determination
labels ‘adspersus teste Cand., Chota Nagp. Bengale’ and ‘Cand., C.R. Soc. Ent.
Belge 1890, Ann. Soc. Ent. Belge 1892’. Beside the specimen there are two labels,
also in Fleutiaux’s handwriting ‘non adspersus 1857’ and ‘non adspersus Cand.
Mus. Brus. 1928’. The specimen is not conspecific with the lectotype. The IRSNB,
Brussels also possesses a number of specimens standing as adspersus Candéze includ-
ing one bearing Candéze’s label ‘adspersus Cand., Beng. P. Card.’. These specimens
are not conspecific with the lectotype. Whether they are conspecific with the

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 23

specimens in the MNHN, Paris has not been verified. These specimens belong to
one or more species unknown to me.

Adelocera aethiopicus (Candéze) comb. rev.

Lacon aethiopicus Candéze, 1882 : 7.
Lacon lineatus Candéze, 1897: 8. Syn.n.
Adelocera aethiopicus (Candéze) Fleutiaux, 1926 : 96.

L. aethiopicus Candéze. LECTOTYPE (present designation). ETHIopPia: 4,
Bogos, 1870, Sciotel, O. Beccari; n.sp. Aethiopicus Cdz. Abyssinia [Candéze, yellow
border]; Collection E. Candéze; Lacon aethiopicus Cand., det E. Candéze [IRSNB
curatorial label]. (IRSNB, Brussels). The specimen measures 6 mm, compared
with the published length of 5 mm.

The material recorded by Fleutiaux (1919 : 17) from Taveta, 2 g, 2 9 (BMNH),
8 J, 11 2, (MNHN, Paris), Pori de Serengeti, 1 9, (MNHN, Paris) and (1922 : 589)
Sud du Lac Rudolphe, 3 § (MNHN, Paris) is not conspecific and belongs to an
unknown species. Cobos (1964 : 586) records a single female from Ennedi. This
specimen, which is in the Cobos collection, is not aethiopicus Candéze. I believe
it may be testaceus Fleutiaux, though the female of this species is unknown to me.

L. lineatus Candéze. LECTOTYPE (present designation). ETHiopiaA: 2, Choa
[Shoa]; C. Fairm.; lineatus Cand. Abyssinie [Candéze, yellow border]; Collection E.
Candéze; Lacon lineatus Cd., det. E. Candéze [IRSNB curatorial label] (IRSNB,
Brussels).

Other material examined. SOUTHERN YEMEN: Dhala, 4800 ft, 14.ix.1937,
Ig (BMNH). Somati Repustic: Hargeisia, 23-28.6.1963, 3 ¢ (BMNH); Duruksi,
20.v.1961, I g, 1 9 (BMNH); Wardair, 6°56’N., 45°20’E. viii.1945, open high bush
with scattered grass, 19 (BMNH). Kenya: Turkana Prov., 2 9 (BMNH).

Adelocera alluaudi Fleutiaux comb. rev.

Adelocera alluaudi Fleutiaux, 1919 : 9.
Lacon alluaudi (Fleutiaux) Fleutiaux, 1926 : 96.

_ LECTOTYPE (present designation). Tanzania: dg, Afr. or. allemande,
| Kilimandjaro, versant sud-est, Alluaud & Jeannel; Zone inferieure, Riviere Himo,
1000 m, mars 1912, St.66; 2° et 3° articles des antennes petits, egaux [Fleut.];
Adelocera alluaudi Fleut., type [Fleut.]. (MNHN, Paris).

Paralectotype. TANZANIA: 3, Afr. or. allemande, Neu Moshi (alt. 800 m) avril
1912, st. 72; Adelocera alluaudi Fleut., co-type [Fleut.] (MNHN, Paris).

Adelocera andrewsi (Fleutiaux) comb. n.
| Agraeus andvewsi Fleutiaux, 19354 : 23.

|
| LECTOTYPE (present designation). Burma: 9, Burmah; Tharrawaddy, C. Q.

|

24 C. M. F. von HAYEK

Corbett, coll. Andrewes; andrewsi Fleut., type [Fleut.]; andrewsi Fleut., n.sp.
[Fleut.] (MNHN, Paris).
Paralectotype. Burma: 9, Tharrawaddy, Burma; Agraeus andrewsi Fleut.
[Fleut.] 1 ex., Pegu [17°20’N., 65°29’E.] India (MNHN, Paris). Abdomen missing.
I have not had the opportunity of examining the syntype material from Katha
[24°r0'N., 96°21’E.] (FRI, Dehra Dun).

Adelocera baccatus (Fleutiaux) comb. n.

Agraeus baccatus Fleutiaux, 1935a : 26; figs.

LECTOTYPE (present designation). Java: 9, Java; Agraeus baccatus Fleut.,
Type [Fleut.]. (MNHN, Paris).

Paralectotype. JAVA: 9, Java; communiqué a Bléte, Mus. de Leyde [Fleut.].
(MNHN, Paris).

Dr Wiebes (in litt.) reports that the specimen recorded by Fleutiaux from Java,
Wonosoba, mai (E. Jacobsen) cannot be found in the RNH, Leyden.

Adelocera bakeri (Fleutiaux) comb. n.

Tvachylacon bakeri Fleutiaux, 1935a : 14.

Holotype. PHILIPPINES: 9, Philipines, Imugin, N. Viscaya, Baker; Adelocera
bakeri Fleut., type [Fleut.] (MNHN, Paris).

Adelocera beauchenei (Fleutiaux) comb. rev.

Lacon beauchenei Fleutiaux, 1918d : 195.

Adelocera beauchenei (Fleutiaux) Fleutiaux, 1926 : 96.

Brachylacon beauchenei (Fleutiaux) Fleutiaux, 1927 : 90; fig. 1.
Brachylacon beauchenei (Fleutiaux); Fleutiaux, 1947 : 254.
Brachylacon (Aganolacon) beauchenei (Fleutiaux); Ohira, 1967a@ : 55.

LECTOTYPE (present designation). NortH VIETNAM: 4, Bao Lac, Tonkin;
Museum Paris, Coll. E. Fleutiaux; Lacon beaucheni Fleut. type: Ann. Soc. Ent.
Fr. 1918 : 195. Collection Fleutiaux [Fleut.] (MNHN, Paris).

Other material examined. NorTH VIETNAM: Hoa Binh, Lac Tho, I ex. (Fleutiaux,
1927 and 1947 locality). (MNHN, Paris).

Adelocera bhasini (Fleutiaux) comb. n.

Agraeus bhasini Fleutiaux, 1932d : 228, figs.
Agvaeus bhasini Fleutiaux; Fleutiaux, 19354 : 28.

Lectotype (designated by Fleutiaux, 1935a). Inp1A: 9, New Forest, Dehra
Dun, G.D. Bhasin, 13.viii [sic].1928; on Litsea polyantha; Agraeus bhasini Fleut.
[Fleut.] (MNHN, Paris).

Fleutiaux based his description on two specimens. I have not had the oppor-

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 25

tunity of examining the paralectotype from Landstowne [sic, perhaps Landsdowne,
29°52'N., 78°41’E.} Maurs, 2500 pieds (M. M. Bhatia) mars (FRI, Dehra, Dun).

Adelocera birmanicus (Fleutiaux) comb. n.

Agvaeus birmanicus Fleutiaux, 1935a : 22.

Holotype. Burma: 4g, Toungoo; Toungoo, Birmanie; Andrews; birmanicus
Fleut., type [Fleut.]; birmanicus n.sp. [Fleut.]; Coll. H. E. Andrewes; 1934.34
(BMNH).

Adelocera borneensis (Fleutiaux) comb. n.

Trachylacon borneensis Fleutiaux, 1935a@ : 15.

LECTOTYPE (present designation). BORNEO: J, 59780 [Fry catalogue number
= South East Borneo, Tamyang Lazary (position unknown). Dr Schreiber,
Germ.Miss.]._ German Mission; Borneo, S. East; Fry coll. 1903.130; Trachylacon
borneensis Fleut., type [Fleut.]; Trachylacon borneensis n.sp. [Fleut.] (BMNH).

Paralectotype. 4, same locality as the lectotype with Fleutiaux’s determination
label and a separate printed ‘co-type’ label (MNHN, Paris).

Adelocera brunneus (Lewis)

Lacon brunneus Lewis, 1894 : 29.
Adelocera brunneus (Lewis) Fleutiaux, 1926 : 96.

Holotype. JAPAN: g, Japan, G. Lewis, 1910.320; Oyama, 24.v—26.v.80; Lacon
brunneus Lewis, type [Lewis] (BMNH).

The specimen from Ceylon recorded by Lewis (loc. cit.) is in the BMNH. It
bears the following labels: 9, Ceylon, G. Lewis, 19g10.320; Bengawantalawa, goo-
5200 ft, 28.11-12.111.82; Lacon brunneus Lewis [Lewis]. As Lewis observes, it is
very similar to the holotype, but in my opinion it is not conspecific with it.

I have not had the opportunity of examining the specimens recorded by Miwa
(1934 : 9) and Ohira (1954) : 14 and 19670 : 105).

Adelocera cambodiensis (Fleutiaux) comb. n.

[Lacon tumens Candéze; Fleutiaux, 1918d : 195. Misidentification.]
Brachylacon cambodiensis Fleutiaux, 1927 : 92.
Brachylacon cambodiensis Fleutiaux; Fleutiaux, 1947 : 253.

LECTOTYPE (present designation). CAmBopiA: gj, Camboge Kampong Thon,
Vitalis; Lacon tumens Cand. Fleut., 1918; Lacon cambodiensis Fleut., type [Fleut].
(MNHN, Paris).

Paralectotype. 1 ex., Laos, Vientiane, May 1915, R. V. de Salvaza; Adelocera
(Brachylacon) cambodiensis Fleut. [Fleut.] (BMNH).

I have not seen the second specimen recorded from the type-locality by Fleutiaux
(19184).

26 Cl Ma ES von HANG

Adelocera catulus (Candéze) comb. n.

Agvaeus catulus Candéze, 18930 : II, ¢.
Agrvaeus catulus Candéze; Fleutiaux, 19354 : 27.

Lectotype (designated by Fleutiaux, 1935a). JAVA: g, Java oriental; Mts
Tengger, Fruhst.; Type: n.sp. iv.gi [sic] catulus Cand , Java or. Fr. [Cand., yellow
border]; Collection E. Candéze; Agraeus catulus Cd. det. E. Candéze [IRSNB
curatorial label]; catulus Cand., Fleut. 1934 rev. [Fleut.] (IRSNB, Brussels).

Candéze based his description on six specimens. Fleutiaux (1935a@ : 27)
designated the lectotype and described the remaining five specimens, which both
he and Candéze believed to be females, as Tvachylacon confusus (see below).

Other material examined. JAvaA: Detr. de la Sonde [?Sunda Straits], 1 J (Fleu-
tiaux, 1935a locality) (MNHN, Paris); Toegoe [position unknown: there are a
number of villages of this name in Java], I g (RNH, Leyden).

Adelocera confusus (Fleutiaux) comb. n.

Trvachylacon confusus Fleutiaux, 19354 : 16.
Agraeus catulus Candéze, 1893b : 11, 9.

Fleutiaux based his description on five specimens which Candéze believed to be
females of A. catulus Candéze.

LECTOTYPE (present designation). JAva: 3, Java orientale; Collection E.
Candéze; Agraeus catulus Cd., det. E. Candéze [IRSNB curatorial label]; Trachy-
lacon confusus Fleut., pour Agraeus catulus 2 Cand., Fleut. rev. 1934 [Fleut.]
(IRSNB, Brussels).

Paralectotypes. I 3, with the same labels as the lectotype, (MNHN, Paris).
I g, 2, Java oriental; Collection E. Candéze; Agraeus catulus Cd. det. E. Candéze
[IRSNB curatorial label]; E. Fleutiaux rev. 1934. Trachylacon confusus Fleut.
[IRSNB curatorial label] (IRSNB, Brussels).

Adelocera constrictus (Ritsema) comb. n.

Agvaeus constrictus Ritsema, 1881 : 29.

[A grvaeus feroculus Candéze; Fleutiaux, 1918d : 188. Misidentification.]

Agvaeus plumatus Fleutiaux, 1927 : 95. [Synonymized with constrictus Ritsema by Fleutiaux,
1935@ : 25.]

Cavicoxum monstrosum Pic, 1928 : 21. Syn. n.

A gvaeus monstruosus (Pic); Fleutiaux, 1931 : 74. [Unjustified emendation. |

Agvaeus monstrosus (Pic); Fleutiaux, 1935a : 28.

Cavicoxum monstrosum Pic; Fleutiaux, 1947 : 243.

Agvaeus constrictus Ritsema; Fleutiaux, 1947 : 245.

Agraeus constrictus Ritsema. Holotype. JAvaA: 3, de Gavére, Batavia; Agraeus
constrictus Rits., type [Rits.] (RNH, Leyden).

Agraeus plumatus Fleutiaux. LECTOTYPE (present designation). Laos: 9,
Tathom, Laos, Vitalis. September 1915; Agraeus plumatus Fleut., type [Fleut.];

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 25,

type de la figure [Fleut.]; constrictus Rits., plumatus Fleut., type de la figure
[Fleut.] (MNHN, Paris). Fleutiaux based his description on the specimens he
had misidentified as A. feroculus Candéze in 1918.

Cavicoxum monstrosum Pic. LECTOTYPE (present designation). Laos: Q,
Laos, Kieng Khong, le 17.1v.1920, P. Vitalis de Salvaza; Cavicoxum monstrosum
n.g., n.sp. [Pic]; Genre Agraeus selon Fleutiaux [Pic]; Agraeus monstrosus Pic,
type, sub Cavicoxum monstrosum [Fleut.] (MNHN, Paris).

Other material examined. JAvA: Agraeus monstruosus [sic] Pic, sub Cavicoxum
compare au type [Fleut.], 1 9 (MNHN, Paris), 1 g, 1 2 (BMNH); G. Oengaran,
vi.1907 (Drescher) 1 2; Djogjakarta (C. J. Louwerens), 1 3, 1 9; Afd. Tjiamis, xi.1g01
(Drescher), 1 3 (ZM, Amsterdam). I have not examined the specimen from Java;
Samarang determined as monstrosum Pic by Fleutiaux (1947).

Adelocera cylindriformis Fleutiaux
Adelocera cylindriformis Fleutiaux, 1935) : 198; pl. 8, fig. 8.

LECTOTYPE (present designation). Krnya: gj, Kenya, Lokitang, Turkana
Nord, 750 m; Mission de l’Omo, C. Arambourg, P. A. Chappuis & R. Jeannel,
1932.33; cylindriformis Fleut., type [Fleut.] (MNHN, Paris).

Paralectotype. Eruiop1a: 3, Ethiopie Merid. Bourie [sic, published locality
is Bourillé] Bord de la Riv. Omo, 600 m; Mission de l’Omo, C. Arambourg, P. A.
Chappuis & R. Jeannel 1932-33. (MNHN, Paris).

Both specimens differ from the description in that the metasternal tarsal grooves
do not attain the posterior lateral angles of the metasternum. The grooves are
curved so that the distal end lies at a point near the lateral margin about two-
thirds of the distance from the anterior margin of the metasternum.

Adelocera decipiens (Boheman)

Lacon decipiens Boheman, 1851 : 417.
Adelocera decipiens (Boheman) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation. SouTH Arrica: 9, Caffraria; J. Wahlb.;
_ Typus: decipiens Boh. [Boh.] (NR, Stockholm).

The published locality is ‘Terra Natalensi’. The absence of a more detailed
locality label is discussed on p. 280. The metasternum is very short compared
with species of similar size and appearance, such as the Indian minusculus (Candéze).
This characteristic is generally associated with a reduction in the size of the wings.
Unfortunately the wings have been destroyed by verdigris.

Adelocera demissus (Candéze)

| Lacon demissus Candéze, 1889 : 74.

_ Lacon demissus Candéze; Candéze, 1890 : 34.
Lacon demissus Candéze; Candéze, 1892c : 484.

| Adelocera demissus (Candéze) Fleutiaux, 1926 : 96.

28 C2. Ms By von HAYEK

LECTOTYPE (present designation). INnp1IA: 3, Bengale; Collection E. Candéze;
Lacon demissus Cdz., det E. Candéze [IRSNB curatorial label] (IRSNB, Brussels).

Standing with the lectotype and paralectotype in the IRSNB are four specimens
from the same locality, one of which bears Candéze’s yellow-bordered determination
label. These four specimens differ from the description and lectotype in that the
elytra are distinctly striate. All eight specimens are mounted on cards but have
pin-holes in the right elytra showing that they have been remounted. Since the
elytral puncturation is clearly visible with the naked eye it is improbable that
Candéze included these four in his original series. I believe that Candéze’s
determination label was replaced on the wrong specimen when the material
was remounted. For this reason the general practice of designating the specimen
bearing Candéze’s determination label has not been followed.

Other material examined. InpD1IA: Chota Nagpur, 1 9, determined by Candéze
(Candéze, 1890 and 1892¢ locality) (MNHN, Paris).

Adelocera difficilis (Lewis) sp. rev., comb. n.

Lacon difficilis Lewis, 1894 : 29.
[Lacon trifasciatus Candéze; Candéze, 1873 : 1. Misidentification.]
[Brachylacon microcephalus Motschulsky; Kishii, 1961 : 25, pl. 2, figs 9, 10. Misidentification.]

LECTOTYPE (present designation). JAPAN: Sex undetermined, left hand of
two specimens mounted on one card. 9.4.81 [on underside of card mount]; Japan,
G. Lewis, 1910.320; Nagasaki, 13.ii-21.iv.81; Lacon difficilis Lewis, type [Lewis]
(BMNH).

Paralectotypes: I ex., on card beside lectotype. 490on one card, Nag. (on under-
side of card mount); Japan, G. Lewis, 1910.320. Nagasaki, 13.11.—21.iv.81; Lacon
difficilis Lewis [Lewis] (BMNH).

Other material examined. JAPAN: Janson collection from Lewis, determined
as trifasciatus by Candéze, 1 ex, (BMNH). This is probably part of the series
misidentified by Candéze in 1873 and could be part of Lewis’ syntype series of
difficilis. I have not examined Kishii’s material. The identification is based on
the good figure of the aedeagus.

Fleutiaux (1918d : 194) treated difficilis Lewis as a synonym of Lacon mucro-
cephalus Motschulsky, but examination of the type-material of the two species
has shown that he was mistaken. Up to the present time I have not seen any
difficilis specimens from localities other than Japan, while microcephalus appears
to be restricted to Ceylon (see p. 38).

Kishii (1961 : 25), Miwa (1933 : 29, 1934 : 70, 85) and Ohira (1934 : 10, 1968 : 73,
1969: 94 & 1969c: 28) have recorded microcephalus Motschulsky from Japan.
It is probable that re-examination of the material will show it to be difficilis Lewis.

Adelocera dilatatus (Fleutiaux) comb. rev.

Lacon dilatatus Fleutiaux, 1903a@ : 572.
Lacon dilatatus Fleutiaux; Fleutiaux, 19184 : 208.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 29

Adelocera dilatatus (Fleutiaux) Fleutiaux, 1926 : 96.
Brachylacon dilatatus (Fleutiaux) Fleutiaux, 1927 : 90.
Brachylacon dilatatus (Fleutiaux); Fleutiaux, 1947 : 254.

LECTOTYPE (present designation). NortH VIETNAM: Laokay, (Tonkin);
Lacon dilatatus Fleut., type [Fleut.|; L. dilatatus Fleut., type, Ann. Soc. Ent. Fr.
1902 p. 257 [Fleut.] (MNHN, Paris).

Other material examined. NorTH VIETNAM: Lac Tho., 1927, 2 ex., (Fleutiaux,
1927 locality) (MNHN, Paris); Vallee de la Haute Riv. Claire, between Hapiang
[22°49/N., 104°48’E.] and Yikh-Tuy, 1 ¢ (Fleutiaux, 1947 locality) (MNHN, Paris).

Adelocera discedens (Candéze) comb. n.

Pericus discedens Candéze, 1892 : 485.
Pericus discedens Candéze; Fleutiaux, 1905 : 320.

LECTOTYPE (present designation). INDIA: g, Bengale; n.sp. 1892, discedens
Cand., Bengale, Barwai, P. Card. [Cand., yellow border]. Type; Collection E.
Candéze. (IRSNB, Brussels).

Paralectotype. InpIA: I ex., Bengale; Collection E. Candéze Agraeus discedens
det. E. Candéze [IRSNB curatorial label]. (IRSNB, Brussels).

The published type-locality is Sunk [Sank] Valley, July. For notes on the
localities of Cardon’s material see p. 273.

Adelocera doherti (Fleutiaux) comb. n.

Trachylacon doherti Fleutiaux, 1935a@ : 17.

LECTOTYPE (present designation). Inp1A: g, 61856. [Fry coll. catalogue =
Margherita, Patkai Mts., under 2000 ft]. Doherty; Assam, Patkai Mts.; Fry
coll. 1905.100, Trachylacon doherti Fleut., type [Fleut.] (BMNH).

Paralectotypes. InpIA: 1 ex., Doherty: India or., Manipur; Fry Coll. 1905.100
(BMNH). 1 4, Doherty; India or., Assam Valley; Fry Coll. 1905.100. (MNHN,
Paris).

Adelocera excavatus (Fleutiaux) comb. n.

Agraeus excavatus Fleutiaux, 1918a : 207.

Agraeus excavatus Fleutiaux; Fleutiaux, 1927 : 94; fig. and pl. 2, fig. 2.
Agvaeus coomani Fleutiaux, 1927: 97. Syn.n.

Agraeus excavatus Fleutiaux; Fleutiaux, 1931 : 74.

Agraeus coomani Fleutiaux; Fleutiaux, 1935a : 22.

Agraeus excavatus Fleutiaux; Fleutiaux, 19354 : 28.

Agraeus excavatus Fleutiaux; Fleutiaux, 1947 : 246.

Agrvaeus coomani Fleutiaux; Fleutiaux, 1947 : 248.

Xa Rac, Province de Thaudamont, forets de Bambous, Capus; Agraeus excavatus

Agraeus excavatus Fleutiaux. Holotype. SouTH VIETNAM: 9, Cochinechine,
| Fleut., Type [Fleut.] (MNHN, Paris).

30 G. M. Fo von HAYEK

Agraeus coomani Fleutiaux. LECTOTYPE (present designation). Nortu
VIETNAM: 4, Tonkin, Lac Tho, Hoa Binh, A. de Cooman; Type; Agraeus coomani
Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes: 14 3, with same locality as lectotype; 7 ex. bear Fleutiaux’s
determination labels and one an additional label in Fleutiaux’s hand ‘type de la
figure’, (MNHN, Paris). 1 3g, same locality as lectotype, with Fleutiaux’s deter-
mination label (BMNH).

Other material examined. NoRTH VIETNAM: Lac Tho, 5 2? (MNHN, Paris),
I ¢ (BMNH) (Fleutiaux, 1931, locality). Laos: Kieng Khouang, iv.1g64, 2 ¢
(BMNH). Burma: Karen Hills, Thandaung, 1 3 (Fleutiaux, 19352, locality)
(BMNH). Inp1a: Ranikhet Divn. [29°38’N., 79°28’E.] Kumaon, vii. 1g21, 1 ¢
(

Adelocera exiguus (Candéze)

Lacon exiguus Candéze, 1897 : Io.
Adelocera exiguus (Candéze), Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). BORNEO: jg, Bangwey [Banggi] Borneo;
n.sp. 1895, exiguus Cand., 1. Bangwey St. [Cand., yellow border] Lacon exiguus
Cd., det. E. Candéze [IRSNB curatorial label]; Type (IRSNB, Brussels).

Other material examined. BORNEO: Bangwey, 3 ex. (MNHN, Paris).

The lectotype differs from the description in that it measures 3 mm in length
and I mm in width compared with Candéze’s measurements of 1 3/4 mm and 2/3 mm.
In addition the elytra are striate and tarsal grooves are present on the propleurae
and metasternum. However, I have no doubt that this is the material on which
Candéze based his description. His errors are almost certainly due to the very
small size of the specimen.

Adelocera falsus (Fleutiaux) comb. n.
Agraeus falsus Fleutiaux, 1927 : 97, fig. and pl. 2, fig. 27.
[Agraeus mouhoti Candéze; Fleutiaux, 1918d : 188. Misidentification.]
Agraeus falsus Fleutiaux; Fleutiaux, 1947 : 248.

LECTOTYPE (present designation).. NORTH VIETNAM: 4, Tayninh, 6.10;
Capitaine Fouguet; mouhoti Cand., [Fleut., crossed out]; A. falsus Fleut., type,
mouhoti Fleut., non Cand. [Fleut.] (MNHN, Paris).

Paralectotypes. SOUTH VIETNAM: I g, Cochinechine, Mont de Chaudoc, Har-
mand, 1887; Agraeus mouhoti Cand. [Fleut.]; Agraeus falsus Fleut. non Cand.
[Fleut.] (MNHN, Paris). 1 g, Cochinechine, Cap St. Jaques, collection Cordier; A.
falsus n. sp. Blair, Brit. Mus. 1929 [Fleut.]; type de la figure [Fleut.] (MNHN, Paris).

At the present time the female of this species is unknown. By analogy with
species whose males resemble /falsus Fleutiaux (e.g. excavatus Fleutiaux, lobicollis
Motschulsky) it seems probable that the prothorax of the female has strongly
modified anterior angles.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 31

Adelocera feai (Fleutiaux) comb. n.

Agvaeus feai Fleutiaux, 19354 : 23.

[Agraeus mouhoti Candéze; Candéze, 1888 : 671 pars, g not 9 as recorded. Misidentification.
Synonymized by Fleutiaux, 19354 : 23.]

[Agraeus mouhoti Candéze; Candéze, 1891a : 755 pars. Misidentification. Synonymized by
Fleutiaux, 1935¢@ : 23.]

LECTOTYPE (present designation). Burma: g, Maymyo, vi1o. H. L.
Andrewes; Maymyo, Birmanie; Andrewes; feai Fleut., type [Fleut]; feai n.sp.
[Fleut.] (MNHN, Paris).

Paralectotypes. Burma: 1 g, Maymyo v.10. H. L. Andrewes; Agraeus feai
Fleut. [Fleut.]; feai n.sp. mouhoti Cand. 1888 (non 1874) [Fleut.]. (BMNH). 1 J,
Maymyo. v.10. H.L.Andrewes (BMNH). 16, Maymyo,vi.ro. H.L. Andrewes
(BMNH). 1 3, Carin Cheba [see p. 275]. goo-1,000m L. Fea v.xii.88; Agraeus
mouhoti 2 [Cand.] Collection E. Candéze; non mouhoti Cand. 1874 sec Fleut. 1934
[Fleut.]; Agraeus feai Fleut., mouhoti Cand. 1888 non Cand., 1874. 1934 [Fleut.],
(Candéze; 1891 locality) (IRSNB, Brussels). 1 3, Tenasserim, Thagata [position
unknown]. Fea, Apr. 1887; coll. Schwarz; Agraeus mouhoti Cand.; Agraeus
feai Fleut. [Fleut.]. (Candéze; 1888 locality) (DEI, Eberswalde). Inp1A: 1 4,
Gopaldhara, Rungabong Vall. [position unknown] Sikkim, H. Stevens. (BMNH).
I dg, Shillong, Assam, F.W.C.; Agraeus sp. near Lacon [Champ.]; feai Fleut. [Fleut.].
(BMNH). 1 ¢ Assam Valley; Doherty; 61880 [Fry collection catalogue = March
to Kohima from country] Fry coll. 1905.100; Agraeus feai Fleut. [Fleut.] (BMNH).
I g, Assam, Patkai Mts; 61060 [Fry collection catalogue = Patkai Mts, under
2000 ft]; Fry coll. 1905. 100 (BMNH). 1 3g, Assam, Patkai Mts; Doherty; Fry
Coll. 1903. 100 (BMNH). 1 ¢, C. Almora Divn. Kumaon, U.P. July 1921, H.G.C.;
feai Fleut. (Brit. Mus.) [Fleut.] (MNHN, Paris). 4 3, C. Almora Divn Kumaon,
U.P. July ’20,H.G.C. Onespecimen bears Fleutiaux’s determination label. (BMNH).
I g, C. Almora, Kumaon, India. H.G.C. (BMNH). 1 3, Berenang [position
unknown] 6000’, Almora Dt. U.P. R.N. Parker, 30 vi.1923; feai Fleut, Mus. Dehra
Dun [Fleut.] (MNHN, Paris). 1 3g, Kulu [position unknown, possibly Punjab,

| Kangra district 32°09’N., 76°15’E.] 3600 Fuss; Kulu, Himalaya (MNHN, Paris).
| Other material examined. INDIA: Ranikhet, Kumaon, 1 g (MNHN, Paris);
_ Dehra Dun, on grass, I0.vii.I9g12, I ¢ (MNHN, Paris); Dehra Dun, Kaulagarh,
| 73 (MNHN, Paris); Dehra Dun, New Forest, 28.ix.1929, 1 g. (FRI, Dehra Dun).

The identification of Fleutiaux’s syntype material was not a simple task.
Fleutiaux did not record the localities on the labels on the specimens. Instead he
listed Birmanie, Tenasserim, Sikkim, Assam, Almora and Kulu as the regions from
which his material was obtained. Fortunately his collection contains a specimen
| labelled ‘type’ and others with his determination labels and this enabled me to
assemble a syntype series from localities within the regions recorded by Fleutiaux.
Specimens from type-localities but without determination labels have been included

in the syntype series if they stood as feaz in collections mentioned by Fleutiaux.

__ The inclusion of certain specimens from Thagata and Carin Cheba in the syntype-

series requires an explanation. Fleutiaux lists A. mouwhoti Candéze, 1888 : 671

|
|
|

32 Cc. M.. EF. von HAYEK

pars as a synonym of feaz, indicating in a footnote that the material is in the DEI,
Eberswalde (formerly in Berlin-Dahlem). The only specimen in that collection
from Thagata labelled mouhoti (though not by Candéze) and correctly determined
as feai by Fleutiaux is a male. The specimen agrees very well with Candéze’s
description of mouhoti 2 and is assumed to be one of the three specimens (misidenti-
fied as I 9, 2 g) recorded by Candéze (1888). For the other two specimens (2 9)
see lobicollis Motschulsky, other material examined, p. 36.

Fleutiaux’s footnote also refers to A. mouhoti material (number of specimens not
recorded) from Carin Cheba, 9 a 1,100 m, mai a decembre (Fea) apparently also
recorded by Candéze, 1888 in the IRSNB, Brussels. Candéze did not record
mouhoti from Carin Cheba until 1891 (1891a@; 775, 3 9). A male from this locality
in the IRSNB, Brussels labeled Agvaeus mouhoti 9 by Candéze has been correctly
identified as A. feat g by Fleutiaux. The remaining two specimens (?MCSN,
Genoa) have not been examined. As Candéze appears to have misidentified the
sexes it seems probable that these two specimens will be found also to be male
fea Fleutiaux.

Adelocera feroculus (Candéze) comb. n.

Agraeus fevoculus Candéze, 1874 : 105.
Agraeus feroculus Candéze; Fleutiaux, 1935a@ : 25.

Lectotype (designated by Fleutiaux, 1935a). SUMATRA: g, C.W., Sumatra;

Sum.; Janson coll., 1903 : 130; ex Coll. Wallace [Janson]; Agraeus feroculus Candéze,
type [Janson]; feroculus Cdz., type [Cand.] (BMNH).

Adelocera fleutiauxi nom. n.

Prolacon alluaudi Fleutiaux, 1934d : 197.
Adelocera alluaudi (Fleutiaux) comb. n. [Junior secondary homonym of Adeloceva alluaudi
Fleutiaux, 1919.]

LECTOTYPE (present designation). Ivory Coast: g, Assinie Cote occid.
Afrique, Ch. Alluaud, 1886: Prolacon alluaudi Fleut., type [Fleut] (MNHN, Paris).

Paralectotypes. ZAIRE: I g, Museum Paris, Congo Belge Central, Prov. de
Maniema, Kindu, L. Burgeon 1917; Prolacon alluaudi Fleut. [Fleut.] (MNHN,
Paris). CAMEROUN: I 9, Museum Paris, Cameroun, vers le Nord. Dr. Gromier,
1917 (MNHN, Paris).

This species, the type-species of Prolacon, is discussed on p. 16.

Adelocera fryi (Fleutiaux) comb. n.
Agraeus fryi Fleutiaux, 1935a : 28.
Holotype. Inna: 9, 61053 [Fry coll. catalogue = Assam, Margherita, Patkai
Mts, under 2000 ft]; Doherty, Assam, Patkai Mts.; Fry coll. 1905.100; Agraeus

fryi Fleut., type [Fleut] (BMNH).
Fleutiaux (1942 : 3) records this species from North East Burma, Sadon. The NR.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 33

Stockholm possesses a female from this locality bearing Fleutiaux’ determination
label ‘Agraeus fryi Fleut.? type unique in Brit. Mus.’ It is not conspecific with
the lectotype, lacking the long setae on the elytra. It belongs to a species unknown
to me.

Adelocera fulvicollis (Motschulsky) comb. n.

Trochylacon [sic] fulvicollis Motschulsky, 1858 : 61.

Agvaeus fulvicollis (Motschulsky) Schwarz, 1906 : 28.

[Pericus varviegatus Schwarz; Fleutiaux, 1918a : 207. Misidentification.]
[Pericus variegatus Schwarz; Fleutiaux, 1918) : 188. Misidentification.]
[Trachylacon variegatus (Schwarz) Fleutiaux, 1927 : 89. Misidentification. }
Trachylacon distinctus Fleutiaux, 1935a@: 14. Syn. n.

Trachylacon distinctus Fleutiaux; Fleutiaux, 1947 : 250.

Traychylacon fulvicollis Motschulsky. LECTOTYPE (present designation).
6, yellow circle with illegible manuscript; Trachylacon fulvicollis Motsch., Ind.
or.; Trachylacon fulvicollis Motsch. Sept. 1932, Fleutiaux vid [Fleut.] (ZMU,
Moscow).

Paralectotype. d, on same card as the lectotype (ZMU, Moscow). The
specimens are mounted, line ahead, on one card. The specimen furthest from the
pin has been selected as the lectotype.

Motschulsky does not record the locality of this species. It is probably part
of the material from the ‘continent indien’ to which he refers in the paragraph
preceding the description.

Trachylacon distinctus Fleutiaux. LECTOTYPE (present designation). Norrtu
ViETNAM: J, Hoa Binh, P. Cooman; Hoa Binh, Tonkin, de Cooman; Trachylacon
distinctus Fleut., type [Fleut.] (MNHN, Paris). Fleutiaux records the type locality
as Hoa Binh, Lac Tho. His series (see below) includes specimens without deter-

mination labels from this locality. However as this specimen agrees so well with
the description and bears his type label, I have assumed that the discrepancy
between the published locality and the label is due to an oversight by Fleutiaux.

_ Paralectotypes NorRTH VIETNAM: 4 dg, 2 9, Tonkin, Lac Tho, Hoa Binh, A. de
Cooman. 1 4, Choganh, Tonkin, 4. 1916; Tonkin, Cho Ganh, L. Dupont; Pericus
-variegatus Schw. 1918 [Fleut.]. Trachylacon distinctus Fleut. [Fleut.]. 1 4,
Museum Paris, Lachon [sic, published locality Lakhon] Harmand, 1878; Pericus
variegatus ? g§ Schwarz, 1917 [Fleut.] Trachylacon distinctus Fleut., Ponctuation
d’elytres [remainder illegible] [Fleut.]. 1 g, Museum Paris, Tonkin Central, Env.
de Tuyen Quan, A. Weiss, 1901; Pericus variegatus Schw. 9? Fleut. Bull. Mus.
1918 [Fleut.]; distinctus Fleut. [Fleut]. 1 9, Reg. de Luc Nam (Tonkin), L. Blaise;
Coll. Ph. Francois, Coll. L. Bedel; Trachylacon variegatus Schw., Fleut., 1918,
| 1927 [Fleut.]; distinctus Fleut. 1935 [Fleut.]. 2 9, same locality without deter-
| mination labels (MNHN, Paris).
| Other material examined. NortH VIETNAM: Thakek, 1 6 (MNHN, Paris);
Hoa Binh, viii.tg18, 1 g, (BMNH). Hone Konc; 1 ¢ (BMNH). Burma: 1 ¢
| (BMNH).

|
|

34 Cc. M. F. von HAYEK

Adelocera gibberosus (Fleutiaux) comb. n.

Agraeus gibberosus Fleutiaux, 1935a : 29.

Holotype. Matraya: 9, Malaya, Kuala Lumpur, 14 Mile, S. Buloh, May 25th,
1932. H. M. Pendlebury, Ex. Coll. F.M.S. Museum; Brit. Mus. 1934 : 80; Agraeus
sp. G. E. Bryant det; Agraeus gibberosus Fleut., type [Fleut.] (BMNH).

Adelocera hilaris (Candéze)

Lacon hilaris Candéze, 1891a : 744.
Adelocera hilaris (Candéze) Fleutiaux, 1926 : 96.

The description is based on a single specimen (MCSN, Genoa) collected in
Carin Cheba (see p. 275) in December 1888 by Fea.

The inclusion of hilaris Candéze in Adelocera is tentative, based on Candéze’s
comment that it is “comme le precedent’ which is trifasciatus Candéze (= Adelocera
microcephalus Motschulsky, see p. 38).

Adelocera incisus (Fleutiaux) comb. n.

Brachylacon incisus Fleutiaux, 1940b : 32.
Brachylacon incisus Fleutiaux; Lesne, 1940 : 36.

LECTOTYPE (present designation). BOoRNEo: 9, Sandakan, Borneo, Baker;
Brachylacon incisus Fleut., type [Fleut.] (MNHN, Paris).

The lectotype differs from the description in that the prosternal process is not
grooved, though there is a slight depression between the anterior coxae. The
metepisterna are not parallel-sided but triangular and not visible beyond a point
just posterior to the distal end of the metasternal tarsal groove. It would appear
that Fleutiaux misinterpreted the longitudinal carinae on the epipleurae as the
suture between the metepisterna and the epipleurae (see also P. alluaudi, p. 16).

Adelocera indicus (Fleutiaux) comb. n.

Agvaeus indicus Fleutiaux, 1935a : 22.

LECTOTYPE (present designation). Inp1A: 3, Nilgiri Hills; 1280; Andrewes,
indicus Fleut., type [Fleut.], indicus n.sp. [Fleut.] (MNHN, Paris).

Paralectotypes. Inp1a: I 3, Nilgiri Hills, H.L. [sic] Andrewes, Agraeus indicus
Fleut. [Fleut.] (BMNH). 1 3, Nilgiri Hills (BMNH).

Fleutiaux records the locality as ‘Nilgiri Hills, Ouchterlony Valley, 3-500 ft,
fevrier et aout, en battent les abres (H. C. Andrewes), coll. H. E. Andrewes.’ I
have not been able to discover a specimen bearing this information. It is possible
that the number 1280 on the lectotype refers to a list of localities and biological
data which is now lost.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 35

Adelocera kilimandjarensis (Fleutiaux)

Lacon hilimandjarensis Fleutiaux, 1919 : 17.
Adeloceva kilimandjarensis (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). TANzaANnIA: ¢g, Afrique or. allemande,
Kilimandjaro versant sud-est, Alluaud & Jeannel; Zone inferieure, Neu Moschi
800m. Avril, 1912, st.72; Lacon kilimandjarensis Fleut. type [Fleut.] (MNHN,
Paris).

Paralectotypes. 11 ex., same locality as lectotype, 3 of these bear Fleutiaux’s
determination labels and one is in addition marked ‘Co-type’ on the determination
label (MNHN, Paris). 1 3, same locality as lectotype with Fleutiaux’s determina-
tion label (BMNH).

The specimen, a male from Shimoni, regarded as a local form by Fleutiaux is
in the MNHN, Paris. It is correctly identified.

Adelocera laxus (Candéze)

Lacon laxus Candéze, 1874 : 85.
Adelocerva laxus (Candéze) Fleutiaux, 1926 : 96.

Holotype. NeEw GuINnEA: 9, Mysole [Misodl]; laxus Cdze, type [Cand.]; Lacon
laxus Cand., type Mysol [Janson] (BMNH).

The BMNH also possesses the second specimen, a male, tentatively associated
with this species by Candéze. It is impossible at the present time to decide whether
these specimens are the male and female of the same species.

Adelocera lobicollis (Motschulsky) comb. n.

Trachylacon lobicollis Motschulsky, 1858 : 61.

Agraeus mouhoti Candéze, 1874 : 106. [Synonymized by Fleutiaux, 1932e : 79.]

[Agraeus mouhoti Candéze; Candéze, 1888 : 671, pars. Misidentification.]

Agvaeus lobicollis (Motschulsky) Schwarz, 1906 : 28.

Agvaeus duporti Fleutiaux, 1918d: 187. [Synonymized with mouwhoti Candéze, Fleutiaux,
1926 : I00.]

Agvaeus tonkinensis Fleutiaux, 1927: 94. Syn. n.

Agvaeus mouhoti Candéze; Fleutiaux, 1927 : 96; pl. 2. fig. 40.

Agvaeus lobicollis (Motschulsky); Fleutiaux, 1932e : 79.

Agraeus lobicollis (Motschulsky); Fleutiaux, 1935a : 21.

| Agvaeus tonkinensis Fleutiaux; Fleutiaux, 19354a : 27.

Agvaeus tonkinensis Fleutiaux; Fleutiaux, 1947 : 246.
Agraeus lobicollis (Motschulsky); Fleutiaux, 1947 : 247.

Trachylacon lobicollis Motschulsky. LECTOTYPE (present designation). 4d,
| Ind. or.; Trachylacon lobicollis Motsch. Ind. or. [Motch]; Sept. 1932, Fleutiaux
det. [Fleut] (ZMU, Moscow).

: Paralectotype: 3, yellow paper circle; Trachylacon lobicollis Motsch. Ind. or.;
: Sept. 1932, Fleutiaux det. [Fleut] (ZMU, Moscow).

The country of origin is unknown. It is probably Burma, Malaya, Thailand
2 or Indochina.
| Agraeus mouhoti Candéze. LECTOTYPE (present designation). THAILAND:

|
|

36 Cc. M. F. von HAYEK

3, Siam, Pachebon [sic] Muhot; Janson coll. 1903. 130; A. muhoti [sic] Cdze, type
[Cand.]; Agraeus muhoti [sic] Cdze, type [Janson] (BMNH).

Paralectotype. 3, Pachebon, Muhot; Janson coll. 1903.130; Agraeus mouhoti
[Cand]; Agraeus muhoti [sic] Cdze., Cand [Janson]; lobicollis Mot. mouhoti Cand,
1874 duporti Fleut. [Fleut.] (BMNH).

The published locality is Pechabury [? Phet Buri]. I have not been able to
trace the specimens recorded from the Saunders collection (see p. 279).

Agraeus duporti Fleutiaux. LECTOTYPE (present designation). Nortu VIET-
NAM: g, Cho-Ganh, 5-14; Tonkin, Cho-Ganh, L. Duport; Agraeus duporti, type
[Fleut.] (MNHN, Paris).

Paralectotypes. NORTH VIETNAM: 4 J, Tonkin, Cho Ganh, L. Duport; Agraeus
duporti Fleut. [Fleut.]. 2 3, Tonkin, Cho Ganh, L. Duport. Agraeus duporti
Fleu. [Fleut.]; Juin. 1 g Cho Ganh, L. Duport. Juin 6 J, Tonkin, Cho Gank, L.
Duport (MNHN, Paris) 2 3, Tonkin, Cho Ganh, L. Duport; Agraeus duporti
Fleut. [Fleut.] (BMNH).

Agraeus tonkinensis Fleutiaux. LECTOTYPE (present designation). NortH
ViETNAM: 9, Tonkin, Cho Ganh, L. Duport; Agraeus tonkinensis Fleut; type [Fleut.]
(MNHN, Paris).

Paralectotypes. NorTH VIETNAM: I 9, Tonkin; Cho Ganh, L. Duport; tonkinensis
Fleut. [Fleut.]; type de la figure [Fleut.] (MNHN, Paris). 1 9, Tonkin, Hoa-Binh.
August 1918. R. V. de Salvaza; Indo-China, R. V. de Salvaza 1918. 1; Agraeus
tonkinensis Fleut. [Fleut.] (BMNH). 1 9, Luc Nam, Blaise; Agraeus tonkinensis
Fleut. [Fleut.] (MNHN, Paris).

Every known specimen determined as tonkinensis Fleutiaux has been found to
be female. The belief that tonkinensis is the female of lobicollis is based on the
series from Lac Thé (see below) which consists of males corresponding with lobicollis
and females corresponding to tonkinensis.

Other material examined. BuRMA: I 9, Tenasserim, Thagata, Fea, April, 1887;
agraeus mouhoti; Agraeus tonkinensis Fleut. [Fleut.] (Fleutiaux, 1935a@: 27 and
Candéze, 1888 : 671 locality) (MCSN, Genoa). 1 9, Tenasserim, Thagata, Fea,
April, 1887, Agraeus mouhoti Cand.; mouhoti Cdz. Tenasserim [Cand, yellow
border]; Collection E. Candeze; mouhoti Cand. type Ann. Gen. 1888 [Fleut.];
Agraeus lobicollis Mots., mouhoti Cand., duporti Cand. [Fleut.] mouhoti Cand.,
type revis. ex. coll. Janson in Brit. Mus. Siam [Fleut.] (Candéze, 1888 : 671, locality)
(IRSNB, Brussels). NoRTH VEITNAM: Hoa Binh, Lac Théo, 62 J, 20 2 (MNHN,
Paris); 6 ¢ (BMNH); 3 ¢ (DEI, Eberswalde); N. Annam, Prov. Thanh Hoa, v. 42,
I ¢ (MNHN, Paris); Cochinchina, Laithe Bencat, v. 45, 3 ¢ (MNHN, Paris); Lakhon,
2 ¢ (MNHN, Paris). THAILAND: 17.vi.1936, I ¢ (MNHN, Paris).

Candéze (18914 : 775) recorded 3 ¢ collected by Fea in Carin Cheba, 9 4 1,100m,
mai a décembre. One specimen from this locality labelled Agraeus mouhoti 2 by
Candéze has been located in the ISRNB, Brussels. It has been correctly identified
as A. feai Fleutiaux g and is a paralectotype of that species (see p. 31). The
remaining two specimens (? MCSN, Genoa) have not been examined.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 37

Adelocera lucasseni (Candéze) comb. n.

Agraeus lucasseni Candéze, 1893a : 125.
Tvachylacon lucasseni (Candéze) Fleutiaux, 1927 : 88. [Tentative transference. |
Agyvaeus lucasseni Candéze; Fleutiaux, 1935a@ : 25; fig.

Holotype. Java: 3g, Mr. Th. Lucassen, Simpar, 3000 v.Tegal. Java 10, 1890;
Type; Agraeus lucasseni Cand. n.sp.g [Cand.]; Holotype: Van Zwaluwenburg July
1957 (RNH, Leiden).

Adelocera luzonicus (Fleutiaux) comb. n.
Trachylacon luzonicus Fleutiaux, 1935a : 17.

Holotype. PHILIPPINES: 9, Luzon, Mont Banahao; Trachylacon luzonicus Fleut.,
type [Fleut.] (MNHN, Paris).

Adelocera macarthuri (Fleutiaux) comb. n.

Lacon macarthuri Fleutiaux, 1935c : 91.

LECTOTYPE (present designation). Kernya: 3, McArthur, Hola, Tana Riv.,
Nov. 1933; Lacon (Adelocera) macarthuri Fleut., type [Fleut.] (MNHN, Paris).

Other material examined. SOMALI REPUBLIC: Bidi Scionde, Baso Guita, I 9
(MNHN, Paris).

Adelocera maculosus (Candéze) comb. n.

Agraeus maculosus Candéze, 1893a : 125.
Tvachylacon maculosus (Candéze) Fleutiaux, 1927 : 88.
Trachylacon maculosus (Candéze); Fleutiaux, 19354 : 16.

The description is based on a single specimen from Java, Kalibakoeng (Lucassen)
in the RNH, Leiden. The curator, Dr Wiebes, has informed me that it cannot
be found, nor is there any record of the type ever having been in that collection.
Fleutiaux (1935a) states that the type is in the RNH, Leiden but it seems probable

that he mistook the specimen from Palambuan (see below) for the type. Candéze

may have retained the specimen (see p. 271) and if this is so, it should be in the
IRSNB, Brussels.

__ The generic attribution is based on the three specimens recorded below.

| Material examined. JAvA: Java merid., Palambuan, 1892, H. Fruhstorfer;
Agraeus maculosus; Gen. Trachylacon Mots. [Fleut.], 1 g. Samarang, Nov. rIg19,
I g (RNH, Leiden). Sukabumi, 1 g. Tiji Wynkoopsbaai, 1 ¢ (Fleutiaux, 19354,
localities) (MNHN, Paris).

Adelocera mannerheimii (Candéze) comb. n.

| Agvaeus mannerheimii Candéze, 1857 : 166, pl. 3, fig. 4.
Agraeus cuniculus Candéze. 1874: 105. Syn.n.
| Agraeus ritsemai Candéze, 1883 : 204. [Synonymized with mannerheimii Candéze by Fleutiaux,
| 19354 : 26.]
| Agraeus mannerheimi [sic] Candéze; Fleutiaux, 19354 : 26.

38 C. M. F. von HAYEK

Agraeus mannerheimu Candéze. LECTOTYPE (present designation). 9, India
(Janson); Janson Coll. 1903 : 130; ex. coll. Dejean [Janson]; Agraeus mannerheimi,
type Dj. Cdze [Janson] (BMNH).

Candéze records the locality as Java and comments ‘Je ne l’ai vu que dans la
collection de M. de la Ferté Senectére portant le nom que je lui conservé et sous
laquelle il a été donné par M. Boheman au conte Dejean’. (For notes on the
Laferté Senectére and Dejean collections see pp. 276 and 274.) The absence of the
original determination label and the locality ‘India’ is probably due to Janson
(see p. 276) I have not seen any other specimens of this species from India.

Agraeus cuniculus Candéze. LECTOTYPE (present designation). MALAya: 9,

C. W. Singapore [Janson]; Sing.; Janson Coll. 1903 : 130; A. cuniculus Cdze., type
[Cand.]; ex. coll. Wallace; Agraeus cuniculus Cdze. type [Janson] (BMNH).

Agraeus ritsemae Candéze. LECTOTYPE (present designation). JAVA: dg,
Agraeus ritsemae Cdze., type [Cand.]; Hulié, Java (RNH, Leiden).

Candéze believed that ritsemae might be the female of mannerheimit, which is not
surprising since he was unaware of the very unusual form of sexual dimorphism
(see p. 21) shown by this species. Fleutiaux (1935a@ : 26) identified the sexes
correctly.

Other material examined. JAvA: Bogor, 20.xii.1953, I 2 (BMNH). Borneo:
Martapura, 1 g¢ (BMNH). Sumatra: Mirang, 1 g (BMNH). PHILIPPINEs:
N. Palawan, Binaluan, 1 9 (MNHU, Berlin). Matava: Singapore, I g, 6 Q
(BMNH). Kuala Lumpur, iv. 1931, ix. 1932, viii. 1922, 3 ¢ (BMNH); S. Perak,
Kintra Valley, ix, x, 1 9 (BMNH); W. Coast Langkawi Is., iv. 1928, 1 g, 19
(BMNH).

I have not seen the specimen from Sumatra, Serdang in the RNH, Leiden
recorded by Candéze (1883: 205) or the material from Singapore (Biro) in the
TM, Bundapest recorded by Szombathy (1909 : 120).

Adelocera microcephalus (Motschulsky) comb. rev.

Brachylacon microcephalus Motschulsky, 1858 : 60.

Brachylacon microcephalus Motschulsky; Motschulsky, 1861 : 118.

Lacon trifasciatus Candéze, 1865 : 10. [Synonymized by Candéze, 1891Ic : 24.]
Lacon mucrocephalus (Motschulsky) Gemminger & Harold, 1869 : 1993.
Brachylacon microcephalus Motschulsky; Fleutiaux, 1925 : 207.

Adeloceva microcephalus (Motschulsky) Fleutiaux, 1926 : 96.

Brachylacon microcephalus Motschulsky; Fleutiaux, 1947 : 252.

Brachylacon microcephalus Motschulsky. LECTOTYPE (present designation).
CEYLON: Q, grey paper square; yellow circle with illegible writing; Brachylacon
microcephalus Motsch. Mt. Nura Ellie, Ceylon [Motschulsky]; Sept. 1932.
Fleutiaux vid. [Fleut.] (ZMU Moscow).

Paralectotype. 9, head and prothorax lost; bears a grey square and yellow
circle, similar to those on the lectotype (ZMU, Moscow).

Lacon trifasciatus Candéze. LECTOTYPE (present designation). CEYLON: 9,

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 39

Ceylon, Janson coll. 1903 : 130; Lacon trifasciatus Cdz., Ceylon (Cand.); Lacon
trifasciatus Cand., ex coll. Cand. [Janson] (BMNH).

Candéze (1891 : 24) listed microcephalus Motschulsky as a synonym of trifasciatus
Candéze, disregarding the fact that Motschulsky’s description was published seven
years before his own. Fleutiaux (1918d : 194) corrected the synonymy.

Other material examined. CEYLON: I ¢ (ex Fry coll.), 2 2 (ex Janson Coll.);
Dikoya, 3,800-4,200 ft, 21.i-7.11.82 (Lewis), 2 3, one with Lewis’s determination
label: Lacon trifasciatus Cand. (BMNH).

Lacon muicrocephalus Motschulsky has been recorded from many other localities.
The following specimens have been examined and found not to be conspecific with
the lectotype; they belong to a number of different species unknown to me. INDIA:
Chota Nagpur (Candéze, 1892 : 485, as trifasciatus Candéze), 1 2 (determined by
Candéze), 2 § (MNHU, Berlin); 14 ex., one labelled ‘trifasciatus teste Cand.’
(MNHU, Paris); 1 ex. (BMNH); Frazerpet, Coorg (Fleutiaux, 1933 : 3), 1 d (FRI,
Dehra Dun). Burma: Tienzo [position unknown] (Candéze, 1888 : 91, as trifasciatus
Candéze), 1 ex. (MCSN, Genoa). Maraya: Malacca (Fleutiaux, 1918d : 194), 3 2
(MNHN, Paris). Camsopia: (Fleutiaux, 1927 : 92) 9 ex. (MNHN, Paris). Nortu
VIETNAM: Hoa Binh, Lac Tho (Fleutiaux, 1927 : 92), 10 ex. (MNHN, Paris); Region
de Luc Nam (Fleutiaux, loc. cit.), 1 ex. (MNHN, Paris). Borneo: (Candéze,
1892c : 485, as trifasciatus Candéze and Fleutiaux, 1918d : 194) 3 ex. (MNHN, Paris).
PHILIPPINES: (Fleutiaux, 1916: 220, as trifasciatus Candéze, and 1918d : 194)
Luzon, Mt Maquilling, 8 ex. (MNHN, Paris); Mindanao, 1 ex. (MNHN, Paris);
Butan, 1 ex. (MNHN, Paris); Palawan, 1 ex. (MNHN, Paris).

I have not examined the specimens recorded from Sikkim and Bengal (Candéze,
1892 : 485), Burma, Palon (Candéze, 1891a : 777), Sumatra, Si Rambe (Candéze,
1894 : 486), S. Vietnam, Cochinchina (Candéze, 1892: 485), Japan, Formosa,
Korea and Tsushima Island (Miwa, 1933 : 27, 29, 1934: 70, 185, Kishii, 1961 : 25,
Ohira, 1968a : 73, Ohira, 1969¢ : 28).

Up to the present time all the specimens of microcephalus Motschulsky known to
me are from Ceylon. It seems probable that the species is restricted to that island.

.
| Adelocera minusculus (Candéze)

| Lacon minusculus Candéze, 1874 : 68.
Adelocera minusculus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). INnp1A: 3, Bombay: L. minusculus Cdze.,

| type (Cand.]; Janson Coll. 1903: 130; Lacon minusculus Cand., type [Gahan]
(BMNH).

Paralectotype. Inp1A: 2, Bombay; Janson Coll. 1903.130 (BMNH).

Adelocera minutus (Candéze) comb. n.

Dilobitarsus minutus Candéze, 1878a : LII (6).

|
| LECTOTYPE (present designation). Brazit: 3, Bahia, Reed; n.sp. minutus

40 C.-M. OES von HAYEK

Cdz. Br. [Cand.]; Collection E. Candéze; Dilobitarsus minutes det. E. Candéze
[IRSNB curatorial label]; ex typis (IRSNB, Brussels).

Adelocera nanus (Boheman)

Lacon nanus Boheman, 1851 : 420.

Lacon troglodytes Boheman, 1851 : 419. [Synonymized by Candéze, 1857 : 131.]
Lacon nanus Boheman; Candéze, 1874 : 56.

Adeloceva nanus (Boheman) Fleutiaux, 1926 : 96.

Lacon nanus Boheman. LECTOTYPE (present designation). SouTH AFRICA:
3, Caffraria; J. Wahlb.; Type; nanus [Boh.] (NR, Stockholm).

The published locality is ‘Habitat in terra Natalensi’. The localities of Boheman’s
material are discussed on p. 280. The specimen agrees with the description though
it is slightly larger in size, measuring 3:7 mm in length and 1-5 mm in width
compared with Boheman’s measurements of 34 and I mm.

Lacon troglodytes Boheman. LECTOTYPE (present designation). SouTH
AFRrica: 9, Caffraria; J. Wahlb; troglodytes Bhm. Ins. Caffr. [Boh.] (NR, Stockholm).

The published locality is ‘Habitat in tractibus fluvii Gariepis’. The localities
of Boheman’s material are discussed on p. 280. The specimen agrees reasonably
well with the description though it is slightly larger, measuring 4mm in length
and I-5mm in width compared with Boheman’s measurements of 3?mm and
Ii; mm.

I believe these specimens are the male and female of the same species and that
Candéze was fully justified in placing them in synonymy. Candéze used the name
nanus although troglodytes has page-precedence. However his action, as first
reviser, must be accepted.

Other material examined. SoutH Arrica: Zululand, 49 (BMNH). One specimen
bears Candéze’s determination label ‘nanus’ and is probably the ‘jolie varietie’ of
Candéze (1874 : 56).

Adelocera nebulosa (Candéze) comb. rev.

Lacon nebulosus Candéze, 1857 : 115.

Lacon nebulosus Candéze; Candéze, 1874 : 84.
Adelocera nebulosus (Candéze) Fleutiaux, 1926 : 96.
Brachylacon nebulosus (Candéze) Fleutiaux, 1942 : 3.

LECTOTYPE (present designation). CEYLON: g, Ceylon; Janson coll. 1903 : 130;
Lacon nebulosus Cdz. Ceylon [Cand.]; Lacon nebulosus Cdze. (ex coll. Cand.)
[Janson] (BMNH).

Paralectotype. CEYLON: g, Ceylon; Janson coll. 1903.130; Lacon nebulosus
(ex coll. Cand.) [Janson] (BMNH).

The published locality is ‘les regions orientales et boreales de |’Inde’. I have
not found any specimens bearing these localities. Candéze (1874: 84) records
the locality as ‘Hindoustan; Ceylon’ which in my opinion justifies the lectotype
designation.

:

|

;
1

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 41

The BMNH possesses a male from Rangoon with Candéze label ‘nebulosus
affinis’. This is probably the specimen tentatively attributed to the species by
Candéze (1874). It is not conspecific with the lectotype. Candéze (1892c : 485)
records this species as being ‘rare’ in Chota Nagpore. I have not seen any specimens
from this locality.

Other material examined. INnpIA: South Shan States, road 40m East of
Taunggyi, 25.ix—-13.x.34, I 9; Tenasserim, Mekane, 90km East of Moulmein,
200m, 2-8.xi.34, I 2 (NR, Stockholm), 1 2 (MNHN, Paris). Fleutiaux, 1942,
localities.

Adelocera niger (Schwarz) comb. n.
Pericus niger Schwarz, 1902f : 307.

LECTOTYPE (present designation). Burma: g, Rangoon; Coll. Schwarz;
co-typus, niger Schwarz [Schwarz]; genus Brachylacon, Fleutiaux det., 5.33.
[Fleut.] (DEI, Eberswalde).

Paralectotype. Burma: I jg, Rangoon; Coll. Schwarz, co-typus (DEI,
Eberswalde).

Schwarz’s series usually include a specimen bearing a type-label, but I have
not found a specimen of this species marked in this way. By analogy with
specimens of similar appearance such as nitidus Candéze it seems probable that
when a series including both sexes becomes available niger Schwarz will be found
to be the male of twrgescens Candéze.

Adelocera nitidus (Candéze) comb. n.

Pevicus nitidus Candéze, 1857 : 167; pl. 2, fig. 20.
Pericus sanguinolentus Candéze, 1893c : 172. Syn. n.

Pericus nitidus Candéze. LECTOTYPE (present designation). [Nortu INp1a]:
9, N. India [Janson]; Ind. bor.; Janson Coll. 1903.130; Pericus nitidus Cdze, type,
ex. coll. de Laferté [Janson]; Pericus nitidus [Cand., stuck on underside of Janson’s
label] (BMNH).

The description was based on two specimens from ‘des parties septentrionales
des Indes Orientales’ (see p. 271), one in the de la Ferté and the other in the Deyrolle
collection. The paralectotype has not been located. I have been unable to find
any specimens with labels corresponding to the published type-locality. The
discrepancy between the published locality and that on the label is probably due
to Janson (see p. 276).

Pericus sanguinolentus Candéze. Holotype. Inp1A: 3, Belgaum S., sanguino-
lentus Cand., n.sp. 1893, Bombay, And. [Cand.]; Type: Collection Candéze (IRSNB,

| Brussels). The published type locality and date is Belgaum, Sampgaon, Avril.

The synonymy is based on the examination of a series in the BMNH from
Chipurupali [Chipurupalle, 17°34’N., 33°10’E.], comprising both males and females.
The species displays marked sexual dimorphism. The female is larger than the

42 C. M. F. von HAYEK

male with the prothorax and elytra shining and without puntures. The prothorax
and elytra of the male are moderately strongly and closely punctured.

Adelocera oberndorferi (Schwarz) comb. n.

Pericus oberndorferi Schwarz, 1902b : 202.

LECTOTYPE (present designation). Inp1a: g, Madras; Coll. Schwarz; Type;
oberndorferi Schw. [Schwarz]; Pericus nitidus Cand., Fleut. det.5.1933; kann
nicht nitidus sein dessen Flgd. ‘lisses et sans stries’ sind, Muller det. 1940 (DEI,
Eberswalde).

As far as I am aware Fleutiaux did not publish the nitidus-oberndorferi synonymy.
Other material examined. Inp1A: Madras, xi. 1907, 1 ¢ (BMNH).

Adelocera obesus (Candéze)

Lacon obesus Candéze, 1857 : 132; pl. 2, fig. 18.
A delocera obesus (Candéze) Fleutiaux, 1926 : :6.

LECTOTYPE (present designation). CEYLON: 9, Ceylon, Lacon obesus Cdze.
Ceylon [Cand.]; Janson Coll. ex. Candéze, 1903.130; Lacon obesus Cand., Ceylon
ex. coll. Candéze [Gahan] (BMNH).

Paralectotypes.. Cevrons 1 9, Ceylom:... 22 4- -2een S; 6; diamond shaped blue
label; Mus. Zool. Polonicum Warszawa 12/45. WARSAW t ¢, Ceylon eee eS
99; obesus Cand. Ceylon Nr. [unknown mss.]; Mus. Zool Polonicum Warszawa
12/45 (IZPAN, Warsaw).

Candéze based his description on material ‘communiquée par M. Dohrn’. There
is no means of discovering whether Candéze retained the material or whether he
returned some or all of it. For the history of the Dohrn collection see p. 274. Since
neither of the IZPAN, Warsaw specimens bear Candéze’s determination label, I
have selected the specimen in the BMNH which is known to be from the Candéze
collection as the lectotype.

Other material examined: CEyLon: Horawupotana 16.x.24, 2 2; Woodside,
Urugalla, 9.iv.23, 1 9 (BMNH).

Adelocera oblongus (Fleutiaux) comb. n.

Brachylacon oblongus Fleutiaux, 1934e : 365.

LECTOTYPE (present designation). PHILIPPINES: 9, Manilla, v.19. PI;
Brachylacon oblongus Fleut; type [Fleut.]; Brachylacon microcephalus Motsch.

[?Fleut.] (MNHN, Paris).
Adelocera paeninsularis (Fleutiaux) comb. n.

Tvachylacon paeninsularis Fleutiaux 19354a : 15.

Holotype. Matava: ¢, Singapore, Baker; Trachylacon paeninsularis Fleut.,
type [Fleut.] (MNHN, Paris).

———

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 43

Fleutiaux remarked that this is the only species in which the humeral callosity
is carinate. However, he was mistaken: a similar carina occurs in Adelocera
mucrocephalus Motschulsky.

Adelocera parcus (Boheman)

Lacon parcus Boheman, 1851 : 416.

Lacon occidentalis Candéze, 1857 : 130. Syn. n.

Lacon occidentalis Candéze; Gerstaecker, 1871 : 54.

Lacon occidentalis Candéze; Gerstaecker, 1873 : 174.

Lacon signatus Candéze, 1882 : 6. [Synonymized by Fleutiaux, 1919 : 16.]
Lacon occidentalis Candéze; Fleutiaux, 1921 : 6.

Lacon occidentalis Candéze; Fleutiaux, 1922 : 589.

Adelocera occidentalis (Candéze) Fleutiaux, 1927 : 96.

Adeloceva parcus (Boheman) Fleutiaux, 1926 : 99.

Adelocera occidentalis (Candéze) ; Fleutiaux, 19350 : 1098.

Lacon parcus Boheman. LECTOTYPE (present designation). SOUTH AFRICA:
Caffraria; J. Wahlb., Type: parcus Bhm. [Boh.] (NR, Stockholm). The abdomen
is missing.

The published locality is ‘Habitat in regione fluvii Limpopoensis’. Boheman’s
localities are discussed on p. 280. The specimen measures 5°5 mm in length and
2mm in width compared with Boheman’s measurements of 5 mm and 1} mm.

Candéze (1857 : 161 as charcus, corrected 1874 : 59) comments that he did not
find this species among the specimens he received from Stockholm. It would appear
that the species remained unknown to Candéze, as in 1874: 45 it is listed as sf.
incertae sedis at the end of the key.

Lacon occidentalis Candéze. LECTOTYPE (present designation). SENEGAL:
9, Senegal; occidentalis [Cand.]; Collection Chevrolat; occidentalis Cand., type
Mon. p. 130 [Fleut.]; Fleut. Voy. All. et Jeann. Afr. Or. Elat. p. [Fleut.]. The
specimen stands over a blue Chevrolat collection label ‘Lacon occidentalis Cand.
Mon. 1, 130, 48 Senegalia’ (MNHN, Paris).

Paralectotypes. SENEGAL: I 9, Laf. Senegal; occidentalis Cdz. [Cdz.]: Lacon
occidentalis Cdz. Type, e. coll. de Laferté [Janson] (BMNH). 2 9, Laf. Senegal;
Janson Coll., 1903 : 130; Lacon occidentalis Cdz. Cand. e. coll. de Laferté [Janson]
(BMNH).

Lacon signatus Candéze. LECTOTYPE (present designation). TANZANIA: Q,

_ Zanguebar, Mhonda [in manuscript, next word illegible, probably Ouzigoua, see

below] A. Hacquard, Miss. Ap. 1879 —1** trim. 1880; L. signatus [Cand.]; Museum

| Paris 1952, Coll. R. Oberthur (MNHN, Paris).

Paralectotypes. 9, without labels standing beside lectotype. TANZANIA: 2 9,

| Zanguebar, Mhonda, Ouzigoua. A. Hacquard Mis. ap. 1879 1 trim. 1880;
| [printed label], Museum Paris 1952, coll. R. Oberthur. One specimen bears a

determination label: Lacon signatus Cdz. [Cand.] teste Candéze (MNHN, Paris).
The two specimens with printed labels were not standing with the lectotype when
they were discovered in the Oberthur collection, but there can be no doubt that
they are part of the original series.

44 Cc. M. FEF: von (HAY EE

The description is based on an unrecorded number of specimens from Zanguebar.
This name was formerly applied to a strip of the East African coast between the
mouth of the Juba (or Giuba) River (1°00’S., 42°05’E.) and Cape Delgardo
(10°45’S., 40°38’E.). Mhonda (6°07’S., 37°36’E.) lies about 60 miles from the
coast. Candeéze does not record the provenance of his material so it would be
reasonable to assume that it was in his own collection now in the IRSNB, Brussels
(see p. 271). However, there are no specimens from the type locality determined
by Candéze in that institution. It seems probable that the specimens formed
part of a collection, made by Hacquard at Mhonda and exhibited by Oberthur in
1880 (Bull. Soc. ent. Fr. (5) 10 : CXVIII). Oberthur (loc. cit. : CXIX) states that
the Elateridae had been studied by Candéze and included some new species.

Candéze refers to Gerstaecker (Acad. Berlin Bull. 1866 : 54) [?=Mber. K. preuss.
Akad. Wiss. Berlin) commenting that this is the species considered by Gerstaecker
to be L. occidentalis. To the best of my belief Gerstaecker did not publish any
comments on L. occidentalis in 1866. In 1871 he recorded occidentalis from Lake
Jipe [Kenya] and in 1873 repeated the record, remarking that the species was
previously known from Senegal. The origin of Candéze’s reference is unknown
to me.

Other material examined. SENEGAL: Galam [district, capital Bakel], 1 g; Dakar,
2 46, 5 @ (Fleutiaux, 1919, localities) (MNHN, Paris); Bambey, 3 ex (BMNH);
Sebikotane [14°45’N., 17°08E], vii, 2 ex. (BMNH). Matt: Khayes [=Kayes,
14°26’N., 11°28’E.], I 9 ,2 @ (Fleutiaux, 1919, locality) (MNHN, Paris). GUINEA:
Reg. de Kouroussa, 3 2 (MNHN, Paris). Ivory Coast: Bassam, 1 9 (MNHN,
Paris). DAHOMEY: near Porto Novo, 2 g (MNHN, Paris). Nicerta: S.E. Kano,
Azare, 3 ex. (BMNH). Cuapb: Fort Achambault, Ba-Karé (Boungoul) Kien-Kaga
ou Kiemboga, 1 9 (MNHN, Paris). SupDAN: Delami, v 1 ¢ (MNHN, Paris); Juba,
2 § (ZMU, Helsinki). Eruropia: Bourie (believed to be misprint for Bourillé on
R. Omo, 60 m. N. of L. Rudolf, Fleutiaux, 19358, locality), 2 3, 49 (MNHN, Paris);
I g, 1 2 (BMNH); Hargeisia, vi, 3 g (ZMU, Helsinki). UGanpa: Mt Kadan,
Karamoja, iv, 2g,19(BMNH). Kenya: Lamu L., iv, v, 1916, 1 9 (MNHN, Paris);
Mombassa, 3 2 (Fleutiaux, 1919, locality) (MNHN, Paris); Mont Loroghi, 1 g
(Fleutiaux, 1922, locality) (MNHN, Paris); Turkana Desert, Lokitang [sic], I g
(Fleutiaux, 19350, locality) (MNHN, Paris); Pays Taita: Bura, ili, 1912, I Q
(Fleutiaux, 1919, locality) (MNHN, Paris); Lac Jipe (v. d. Decken), 3 3, 1 9
(Gerstaecker, 1871 and 1873, locality) (MNHU, Berlin). Tanzania: Zanguebar,
I ¢ (Fleutiaux, 1919, locality) (MNHN, Paris); Zanzibar, 1 ¢ (MNHN, Paris);
Bagamoyo, I g§ (IRSNB, Brussels); Tanga, 1 9 (DEI, Eberswalde). MOZAMBIQUE:
Zambeze, Caia, ix, 2 ex. (BMNH); Nova Choupanga, near Chemba, vi, I 9, viii,
I g, xii, r 9 (MNHN, Paris); Pungoué [Pungué] Valley, ii, 3 9, iv, 3 9, vi, I Y,
xi, 9, xii, 1 29 (MNHN, Paris); Barne 15 m. E. of Canxixe, i, 1 9? (MNHN, Paris);
Chiramba, i, I 9, ii, r g, 1 9 (MNHN, Paris); Tambara, ii, 1 9. SouTH AFRICA:
Zoutpan. [? = Soutpansberg] xi, 1 9 (MNHN, Paris).

I have not been able to trace the specimens recorded from Bogos, Abyssinia
by Candéze (1874) and Transvaal by Fleutiaux (19350).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 45

Adelocera perraudieri (Fleutiaux) comb. n.

Brachylacon pervaudievi Fleutiaux, 1927 : 90.

[Lacon nebulosus Candéze; Fleutiaux, 1889 : 139. Misidentification.]
[Lacon nebulosus Candéze; Fleutiaux, 1895 : 685. Misidentification.]
Brachylacon perraudieri Fleutiaux; Fleutiaux, 1947 : 252.

LECTOTYPE (present designation). CamBopia: 92, PP [for Pnomh Penh].
Brachylacon perraudieri Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes: 1 ex., PP; L. nebulosus Cand., Pnomh Penh, Camboge, 1889
[Fleut.]. 19, PP, 1 g, Cap St Jaques (MNHN, Paris).

Adelocera planus (Fleutiaux) comb. n.
Hemicleus planus Fleutiaux, 1919 : 13.
LECTOTYPE (present designation). KENyaA: 9, Afrique Orient. Angl.

Kisoumou (Victoria Nyanza) Ch. Alluaud; Sept.; Hemicleus planus Fleut., type
[Fleut.] (MNHN, Paris).

Paralectotype. KeENyA: 9, Afrique Orient. Angl. Nakuro (Rift Valley) Ch.
Alluaud; Dec.; voir art. ant 2+ 3 [Fleut.]; Hemicleus planus Fleut., Co-type
[Fleut.] (MNHN, Paris).

Adelocera pygmaeus (Baudi)

Lacon pygmaeus Baudi, 1871 : 49.
Lacon pygmaeus Baudi; Marseul, 1872 : 343.
Adelocera pygmaeus (Baudi) Fleutiaux, 1926 : 97.

Holotype. 9, without labels (IMZU, Turin).
The published locality is Cyprus.

Other material examined. Turkey: Adana, 2 ¢ (MNHN, Paris); Namrun,
vi, I 9 (Wewalka collection, Vienna). LEBANON: Beyrouth, 1 9 (MNHN, Paris).

| Adelocera quadriguttatus (Candéze)

Lacon quadriguttatus Candéze, 1895¢ : 485.
| Adelocera quadriguttatus (Candéze) Fleutiaux, 1926 : 96.

Holotype. SUMATRA: sex undetermined, Pangherang Pisang, collected by
_ Modigliani between Nov. 1890 and March 1891 (MCSN, Genoa).

I have not had the opportunity of examining this specimen. The generic
j attribution is based on Candéze’s statement that the species closely resembles
L. nebulosus Candéze.

Adelocera rubicundus (Candéze) comb. n.
Pericus rubicundus Candéze, 1874 : 107.

|

Holotype. Sex undetermined, N; Janson Coll. 1903 : 130; Pericus rubicundus

46 Clavie Ee Vi Ne EAN EA

Cdz., type [Cand.]; Pericus rubicundus Cdze. type [Janson] (BMNH). The genitalia
are missing.

The species was described from an unknown locality. Up to the present time
no additional specimens of this species are known to me.

Adelocera santali (Fleutiaux) comb. n.

Trachylacon santali Fleutiaux, 1933 : 5.
Trachylacon santali Fleutiaux; Fleutiaux, 19354 : 17.

Lectotype (designated by Fleutiaux, 1935a). INDIA: 3, 1.1x.30, Aiyur, N. Salem,
F.R.I. Sandal Survey; Plot 18; Trachylacon santali Fleut., type [Fleut.] (MNHN,
Paris).

Paralectotypes. Two examples from the type-locality, v.30, plot 18 and xi.30,
plot 19 should be in the collections of the FRI, Dehra Dun. I have not had the
opportunity of examining these specimens.

Adelocera schonfeldti (Candéze) comb. n.

Agvaeus schonfeldti Candéze, 1893b : II.
Agvaeus schonfeldti Candéze; Fleutiaux, 1935a : 24.
Agvaeus candezei Fleutiaux, 1935a: 24. Syn. n.

Agraeus schonfeldti Candéze. LECTOTYPE (present designation). BORNEO:
©, Borneo; n. sp. iv. 1891, schonfeldti Cand. Borneo; Sch. [Candéze, yellow border];
Collection E. Candéze: Agraeus schonfeldti Cand. det. E. Candéze [IRSNB
curatorial label]; schonfeldti Cand. ¢, 1934 [Fleut.] (IRSNB, Brussels).

Candéze had specimens of both sexes, but mis-identified them. He believed
that the prothorax of the male possessed a large median tubercle, while that of
the female had no tubercle. Fleutiaux (1935a) records that the type is in the
IRSNB, Brussels, but does not mention any other specimens. He comments
that by analogy with other [related] species, Candéze mis-identified the sexes and
that the type is probably a female. Dissection of the lectotype, which agrees
with the description of the male, has shown this to be so. Candéze’s ‘female’
specimens cannot be found in the IRSNB, Brussels.

Agraeus candezer Fleutiaux. LECTOTYPE (present designation). BORNEO:
6, Riam Kanan, Martapoera, Z. O. Borneo, Doherty, VI-VII; Candezei Fleut.
type; ? schonfeldti Cand., 9 [Fleut.] (MNHN, Paris).

Paralectotypes. JAVA: I g, Java (Meuwen Bay), Detr. de la Sonde, Raffray &
Maindron 1878; Agraeus candezei Fleut., schonfeldti 2 Cand. [Fleut.] (MNHN,
Paris). BORNEO: 2 9, Kuching. J. E. A. Lewis, 1919 : 16; one specimen bears
two labels: near feroculus det. K. G. Blair [Blair]; Candezei Fleut. ? schonfeldti
Cand., 9 (MNHN, Paris). SUMATRA: I g, Sumatra, coll. Schwarz; Mouhoti Cand.
[Schwarz]; ? feroculus Cand., ? schonfeldti Cand. 2 (non 9, sed 3) 8.33 [Fleut.]
(DEI, Eberswalde). Matava: 1 3g, Singapore; cuniculus Cand., Singapore Fl.
[Cand.]; Collection Candéze; Agraeus cuniculus Cd. det. Candéze [IRSNB curatorial

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 47

label]; candezei Fleut., schonfeldti Cand., 2 Fleut. rev. 1934 [Fleut.] (IRSNB,
Brussels). This is obviously the specimen to which Fleutiaux (19354 : 24) refers in
his footnote, though it bears no label to show that it came from the Raffray
collection.

Adelocera serricornis (Quedenfeldt)

Lacon servicornis Quendenfeldt, 1886 : 23, pl. 1, figs 13, 13a.
Adeloceva servicornis (Quedenfeldt) Fleutiaux, 1925 : 205.

LECTOTYPE (present designation). 3, Lacon serricornis Odft. Ogo. [? Queden-
feldt]; Ex Museo Quedenfelt; Paris Museum, 1952, R. Oberthur Coll. (MNHN,
Paris).

Quedenfeldt does not record a precise type-locality. The species was described
from an unrecorded number of specimens collected by Major von Mechow between
1878 and 1881 in the course of expedition to Angola and the Quango River.
According to Horn (1936: 170), part of von Mechow’s collection passed via
Quedenfeldt to the NMHU, Berlin and the remainder to Oberthur (now in the
MNHN, Paris). There are no specimens in the MNHU, Berlin.

Quedenfeldt’s figure is misleading. The third antennal segment, though
triangular, is not as large as the fourth.

Adelocera shirozui (Ohira) comb. n.

Brachylacon (Aganolacon) shivozuwi Ohira, 19674a : 55.
Brachylacon (Aganolacon) shivozui subsp. ishigakiensis Ohira, 1967) : 105.

Brachylacon (Aganolacon) shirozuc Ohira. Holotype. Tatwan: 4, Formosa,
Nanzankei [sic, published locality Nanshanchi] 1965-iv.26, T. Shirozui; Brachylacon
(Aganolacon) shirozui Ohira 1967 [Ohira} (Ohira Coll.).

The author believed the type to be a female. Dissection has shown it to be
a male.

Brachylacon (Aganolacon) shirozut subsp. ishigakiensis. Holotype. JAPAN:
9, Ishigaki (Omotodake) 27.vii.1964, H. Koshi (Shibata coll., Osaka. Ohira, in
litt., specimen not examined).

| Adelocera sparsus (Candéze)

Lacon sparsus Candéze, 1874 : 85.
Adelocera sparsus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). JAVA: 9, Java, C.W.; Janson coll. 1903.130;
sparsus Cdz., type [Cand.]; Lacon sparsus Cand., type, Java [Janson] (BMNH).

The specimen recorded from Thagata, Tenasserim (3, MCSN, Genoa) by Candéze
(1888 : 671) is not conspecific with the lectotype and belongs to a species unknown
‘tome. I have not had the opportunity of examining the specimens from Coorg;
‘Frazerpet and Jawalagiri (? FRI, Dehra Dun) recorded by Fleutiaux (1933 : 5).

|

48 CPM. EF. von HAYEK

Adelocera subcostatus (Candéze)

Lacon subcostatus Candéze, 1891a : 774.
Adelocera subcostatus (Candéze) Fleutiaux, 1926 : 96.

Holotype. Burma: Carin Cheba (see p. 275), collected in March, 1888 by Fea.
MCSN, Genoa.

The generic attribution is tentative, based on Candéze’s comment that this
species belongs to the same ‘section’ as trifasciatus Candéze.

Adelocera succinatus (Candéze)

Lacon succinatus Candéze, 1890 : cl.
Lacon succinatus Candéze; Candéze, 1892¢ : 485.
Adelocera succinatus (Candéze) Fleutiaux, 1926 : 207.

LECTOTYPE (present designation). INnpIA: 9, Bengale; succinatus Cdz. Bengal,
P. Card. [Candéze, yellow border]; Lacon succinatus Cd., det E. Candéze [IRSNB
curatorial label]; Collection E. Candéze (IRSNB, Brussels).

Paralectotypes. INDIA: I ex., Bengale; Lacon succinatus Cd. det. E. Candéze
[IRSNB curatorial label]; Collection E. Candéze (IRSNB, Brussels). 1 ex., Chota
Nagpur, Bengale; succinatus Cand. det. [Cand.]; Cand., C.R. Soc. Ent. Belge 1890,
Ann. soc. Ent. Belg 1892 [Fleut.] (MNHN, Paris).

The published type-locality is Chota Nagpur (see p. 273). Experience has shown
that specimens from this locality may bear only ‘Bengale’ labels. The lectotype
agrees well with the description, although it measures Io mm compared with the
published length of 9mm. Despite the absence of detailed locality labels, I have
no doubt that these specimens formed part of Candéze’s original series.

Adelocera tabaci (Fleutiaux)

Lacon tabaci Fleutiaux, 18956 : 167.
Adelocera tabaci (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). SuMATRA: g, Sumatra; Lacon tabaci
Fleut., type [Fleut.]; L. tabaci Fleut., type, Ann. Belge; 1895: 267 |Fleme
(MNHN, Paris).

Adelocera tanganus nom. n.

Hemicleus minutus Schwarz, 1903a : 358.

Lacon minutus (Schwarz) Fleutiaux, 1919 : 20.

Adelocera minutus (Schwarz) Fleutiaux, 1926 : 96. [Junior secondary homonym of Adelocera
minutus (Candéze, 1878a : LI]1).]

LECTOTYPE (present designation). TANZANIA: 9, Tanga; minutus Schw.
[Schwarz]; Coll. Schwarz; Typus (DEI, Eberswalde). The specimen has suffered
considerable damage. The head and right-hand portion of the pronotum are missing.

Other material examined. KENYA: Voi, 4 ex. (MNHN, Paris), 1 9 (DEL

|

|
|
|
|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 49

Eberswalde); Sambourou (Wa-Nyika), I ex. TANZANIA: Zanguebar, I ex.;
Kilimanjaro, versant sud-est, Riviére Himo, I ex. (Fleutiaux, 1919, localities)
(MNHN, Paris).

Adelocera testaceus (Fleutiaux) comb. n.

Brachylacon testaceus Fleutiaux, 1935¢: 92

LECTOTYPE (present designation). KENYA: J, MacArthur, Ziwani, Apr.
1933; Musée de Nairobi; Brachylacon testaceus Fleut. [Fleut.] (NM, Nairobi).

Paralectotype. I ex., same locality-label as lectotype; Brachylacon testaceus
Fleut., type [Fleut.] (MNHN, Paris). The specimen is very badly damaged.
A great part of the prothorax, the elytra and the entire abdomen are missing.
It is for this reason that I have not followed my usual practice of designating the
specimen marked ‘Type’ by Fleutiaux as the lectotype.

The location of Ziwani is probably 3°23’S, 37°48’E in Kenya and not a place
of the same name at 10°22’S, 40°15’E in Tanzania.

The material recorded from Thua River, xi (MacArthur) has not been located.
See also note on this locality under Lanelater acuminatus (Fleutiaux) p. 242.

The female from Ennendi, Chad, identified as Brachylacon aethiopicus by Cobos
(1964 : 586), may belong to this species.

Adelocera tripartitus (Fleutiaux) comb. n.

Agraeus tripartitus Fleutiaux, 1927 : 98; pl. 2, fig. 36.
Agraeus tripartitus Fleutiaux; Fleutiaux, 1935a : 23.
Agraeus tripartitus Fleutiaux; Fleutiaux, 1947 : 29

LECTOTYPE (present designation). NortH VIETNAM: 3g, Tonkin, Lac Tho,
Hoa Binh, A. de Cooman; Agraeus tripartitus Fleut., type [Fleut.]; type de la
figure [Fleut.] (MNHN, Paris).

Paralectotypes: 4 j, same locality as the type. Two specimens bear Fleutiaux’s
determination label and another Fleutiaux’s labels ‘dessiné par Lesne 1932’ and
‘pieces genitales [which are missing] gardés par Lesne’ (MNHN, Paris).

Other material examined. NortH VIETNAM: I ex. Rakam (Fleutiaux, 1947,
locality) (MNHN, Paris). Laos: 1 g, Xieng Khouang, iv (BMNH).

Adelocera tumens (Candéze)

Lacon tumens Candéze, 1873 : 4.
Adeloceva tumens (Candéze) Fleutiaux, 1926 : 96.

Holotype. JAPAN: g, Lacon tumens n. sp. Cdze. [? Lewis]; Japan, G. Lewis,
1910.320 (BMNH). It would appear that Lewis relabelled the specimen on which
Candéze based his description.

I have not had the opportunity of examining the specimens recorded from
Formosa by Miwa (1929 : 232 and 1934: 70).
D

50 C..M. F. von HAYEK

Adelocera tumidipennis (Candéze)

Lacon tumidipennis Candéze, 1878b : LIV (7).
Lacon pusillus Schwarz, 1898 : 186. [Synonymized by Fleutiaux, 1919 : 20.]
Adeloceva tumidipennis (Candéze) Fleutiaux, 1926 : 96.

Lacon tumidipenms Candéze. Lectotype (designated by Fleutiaux, Ig19g : 20).
TANZANIA: Zanzibar, Raffray (MCSN, Genoa). Not examined.

Paralectotypes. TANZANIA: I 9, Zanzibar, Raffray; Museum Paris 1952, Coll. R.
Oberthur; 251; Lacon tumidipennis Cdz. [Cand.]. 1 9, locality and collection as
above, 252 (MNHN, Paris).

Candéze did not record how many specimens he examined. Fleutiaux (1919)
states that the unique type is in the MCSN, Genoa. However since it is known
(see p. 275) that Fea’s duplicate material was acquired by other collectors, I believe
that the inclusion of the MNHN, Paris material in the type-series is fully justified.

Lacon pusillus Schwarz. LECTOTYPE (present designation). TANZANIA: 3,
Usamb., Coll. Schwarz; Typus; pusillus Schw. [Schwarz] (DEI, Eberswalde).

Paralectotypes. TANZANIA: 2 9, I g originally on the same pin as the type, 2 9,
Usamb.; Coll. Schwarz; Typus. 1 9, Usambara; Coll. Schwarz; Typus. 1 9,
Usamb. Weise; Coll. Schwarz; Typus (DEI, Eberswalde).

Adelocera turgescens (Candéze) comb. rev.

Lacon turgescens Candéze, 1874 : 84.

Lacon turgescens Candéze; Fleutiaux, 1918a@ : 195.

A delocera turgescens (Candéze) Fleutiaux, 1926 : 96.

Brachylacon turgescens (Candéze) Fleutiaux, 1927 : 93, pl. 2, fig. 25.
Brachylacon turgescens (Candéze) Fleutiaux, 1947 : 253.

LECTOTYPE (present designation). MaALaya: 92, Penang [Janson]; Janson
Coll. 1903.130; L. turgescens Cdz. type [Cand.] (BMNH).
The published locality, Paulo Penang, is an alternative spelling for Penang.

Other material examined. NorRTH VIETNAM: Keng-Kabao, I ex. (Fleutiaux,
1927, locality) (MNHN, Paris). Savanakhet, vii, 1 ex. (Fleutiaux, 19182, locality)
(MNHN, Paris). CAmBop1A: Kompong Thon, I g, 1 ex. (Fleutiaux, 19182, locality)
(MNHN, Paris).

Adelocera variegatus (Schwarz) sp. rev., comb. n.

Pericus variegatus Schwarz, 1902b : 203.

Pericus variegatus Schwarz, 1906 : 28; pl. 1, fig. 11.

Trachylacon fulvicollis var. variegatus (Schwarz) Fleutiaux, 1932é : 79.
Trachylacon fulvicollis var. variegatus (Schwarz); Fleutiaux, 19354a : 13.

LECTOTYPE (present designation). Burma: 9, Birma; Coll. Schwarz; Typus;
variegatus Schw. [Schwarz]. Trachylacon fulvicollis var. Fleut. det. 5.33 [Fleut.
(DEI, Eberswalde).

Other material examined. NorTH VIETNAM: I, Tonkin, Lac Tho, Hoa Binh,

:

:

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 51

A. de Cooman; Pericus variegatus Schw., comparé au type [Fleut.]; Brachylacon
fulvicollis Motsch., type elytres [next word illegible] au sommet [Fleut.] (Fleutiaux,
19354, locality) (MNHN, Paris).

Adelocera weisei (Schwarz)

Lacon weisei Schwarz, 1898) : 185.

Lacon weisei Schwarz; Schwarz, 1908 : 75.
Lacon weisei Schwarz; Fleutiaux, 1919 : 20.
Adelocera weisei (Schwarz) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). TANZANIA: 2, Usamb. Weise; Coll.
Schwarz; Typus; weisei Schw. [Schwarz] (DEI, Eberswalde).

Paralectotype. TANZANIA: I 9, locality label as lectotype, but without Schwarz’s
determination label (DEI, Eberswalde).

Other material examined. TANZANIA: Kilimandjaro, Kibonoto, iv, I ex.
(MNHN, Paris); Kilimandjaro (Schwarz, 1908, locality) 1 3, 1 2 (DEI, Eberswalde).
Kenya: Taveta, iii, r § (Fleutiaux, 1919, locality) (MNHN, Paris).

SCAPHODERUS Candéze

Scaphoderus Candéze, 1857 : 46. Type-species: Scaphoderus riehliit Candéze, by monotypy.
Bruyantius Fleutiaux, 1925a : 101. Type-species: Bruwyantius capensis Fleutiaux, by monotypy.
Syn. n.

GENERIC DIAGNOsIs. Each claw bearing a group of setae near the base (Text-fig. 11). Tibial
spurs absent (Text-fig. 10). Mesepisternum does not form part of the margin of the mesocoxal
cavity; mesepimeron forms part of margin of the mesocoxal cavity (Text-fig. 3). Second and
third antennal segments subequal, each smaller than the fourth and following segments
(Text-fig. 13). Antennal groove occupying the anterior three-quarters of the prosternopleural
suture and deep enough to accommodate the antennae. Body clothed with scales. Lateral
margin of pronotum explanate, lateral carina present but not attaining the anterior margin.
A band of longer, more closely packed, erect setae present on the lateral margins of the pronotum.
Propleurae and metasternum without grooves or depressions for the accommodation of the
anterior and middle tarsae. Scutellum simple, without carina. Tarsi simple, without ventral
lobes.

HIsTORY OF THE GENUS. The genus was erected for a single species which
differed from Adelocera (sensu auct., nec Latreille, 1829 = Lacon Castelnau of the
present work) in that the second and third antennal segments are short and from
Lacon (sensu auct., nec Castelnau, 1836 = Agrypnus Eschscholtz of the present
work) in that the antennal groove is longer.

NOTES ON THE GENERIC SYNONYMY. Only the paired tubercles on the anterior
margin of the pronotum and the more strongly explanate margins of the prothorax
distinguish Scaphoderus (? only known) from Bruyantius (g only known). Since
the males and females of certain species (e.g. mannerheimit (Candéze)) belonging
to the closely related genus Adelocera differ from one another in the same way
it seems reasonable to assume that the differences which have been used to separate
the genera are no more than secondary sexual characteristics.

52 CaM: FS von HAMEK

DISTRIBUTION. South Africa.

BIOLOGY AND HABITS. Reichensperger (1915: 19) states that viehlii Candéze
may be myrmecophilous. Otherwise nothing is known of the life-history and
habits of the two species included in the genus.

SPECIES INCLUDED IN THE GENUS

Two species are assigned to the genus. It is debatable whether the presence
of a band of closely packed setae on the lateral margin of the pronotum is of
sufficient importance at the generic level to justify the retention of Scaphoderus
as a Separate genus. When additional material becomes available it may become
apparent that these two species should be included in Adelocera.

Scaphoderus capensis (Fleutiaux) comb. n.
Bruyantius capensis Fleutiaux, 1925 : 101.

LECTOTYPE (present designation). SouTH AFRICA: 3, Museum Paris, Colonie
du Cap. Steynsburg. R. Ellenberger 1914; Bruyantius capensis Fleut., type
[Fleut.] (MNHN, Paris).

Paralectotype. SouTH AFRICA: g, same locality label as lectotype, but without
a determination label (MNHN, Paris).

Scaphoderus riehlii Candéze
Scaphoderus riehlit Candéze, 1857 : 46; pl. I, fig. 3.

Holotype. SouTtH AFrRica: 2, Buq. Natal; Janson coll. 1903.130; Scaphoderus
riehlii Cdz. [Cand.]; Cardiophorus singularis Buq. [Buquet]; Scaphoderus riehlii
Cdze, Cand., type (e coll. de Laferté) (Cardiophorus ? singularis Buq. in mus.)
[Janson] (BMNH).

LACON Castelnau

[Adeloceva sensu auct., nec Latreille, 1829. Misinterpretation.]

Lepidotus Stephens, 1830 : 374. Type-species: Elatey varius Olivier, 1790 [=Lacon quercea
(Herbst, 1874)], by subsequent designation (Hyslop, 1921 : 652). [Homonym of Lepidotus
Asso, 1801 : 38 (Pisces), replaced by Zalepia Arnett, 1953.]

Lacon Castelnau, 1836: 11. Type-species: Elatey atomarius Fabricius, 1789 [=Lacon punctatus
(Herbst, 1779)], by subsequent designation (Hyslop, 1921 : 652).

Ocneus Candéze, 1857 : 87. Type-species: Ocneus limbatus Candéze, by monotypy. Syn. n.

Scelisus Candéze, 1863 : 327. Type-species: Scelisus sanguineus Candéze, by monotypy.
Syn. n.

Alaotypus Schwarz, 1902 : 307. Type-species: Alaotypus subpectinatus Schwarz, by subsequent
designation (Hyslop, 1921 : 625). [Synonymized with Adelocera sensu auct., by Fleutiaux,
t9o18d : 183.]

Sulcilacon Fleutiaux, 1927: 65. Type-species: Adelocera geographica Candéze, by original
designation. Syn. n.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 53

Diphyaulon Arnett, 1952 : 112 [as a subgenus of Lepidotus Stephens]. Type-species: Adelocera
pyrsolepis LeConte, by original designation. [Raised to generic status, Arnett, 1969 : 11.]
Syn. n.

Aulacon Arnett, 1952 : 112 [as a subgenus of Lepidotus Stephens]. Type-species: Adelocera
nobilis Fall, by original designation. [Raised to generic status, Arnett, 1969: 11.) Syn.n.

Zalepia Arnett, 1953: 7. [Replacement name for Lepidotus Stephens; as a subgenus of Lacon
Castelnau. MJaised to generic status, Arnett, 1969: 11.] Syn.n.

KKobulacon Chujo & Ohira, 1965 :2 [as a subgenus of Lacon Castelnau]. Type-species:
Lacon quadrinodatus Lewis, by monotypy. Syn. n.

Lepidelatey Smith, 1969: 11. Type-species: Lepidelater misticius Mignot, by monotypy.
Syn. n.

Arnettia Golbach, 1969 : 155. Type-secies: Adelocera aberrans Candéze, bymonotypy. Syn.n.

Monocyrton Golbach, 1969 : 156 [as a subgenus of Lacon]. Type-species: Adelocera chabannei
Guérin, by original designation. Syn. n.

Cornilacon Golbach, 1969 : 158 [as a subgenus of Lacon]. Type-species: Adelocera longicornis
Champion, by original designation. Syn. n.

Latilacon Golbach, 1969: 158 [as a subgenus of Lacon]. Type-species: Adelocera laticollis
Candéze, by original designation. Syn.n.

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae near the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepimeron forms part of margin of mesocoxal cavity;
metepisternum does not form part of margin of mesocoxal cavity (Text-fig. 3). Second and
third antennal segments not subequal, the third resembles the fourth in shape though it may be
slightly smaller in size (Text-fig. 14). Antennal groove of variable depth, extending at least
half the length of the prosternopleural suture and in many species attaining the anterior coxae.
Body clothed with narrow scales or setae. Prothorax not constricted behind anterior angles;
lateral carinae present. Propleurae with or without grooves or depressions for the accommo-
dation of the anterior tarsi. Metasternum with or without depressions for the accommodation
of the middle tarsi, but never with well defined grooves. Scutellum simple, without longitudinal
carina. Tarsi simple, without ventral lobes.

RANGE OF VARIATION FOUND WITHIN THE GENUS. Species possessing this
combination of diagnostic characters display a wide range of variation in the
appearance of the antennae and the surface of the pronotum and elytra, the type
of body vestiture (scales or setae), its colour and distribution, the length and depth
of the antennal and tarsal grooves or depressions and the appearance of the
mesosternal groove.

Up to the present time these characters have been used, either singly or in
combination with one another, as the diagnostic features of a number of genera
and subgenera. While these may appear justified in works dealing with restricted
faunas, examination of material from all parts of the world has shown the existence
of so many intermediate forms that it is clear that there is no justification for the
retention of the 18 genera and 4 subgenera which are here newly placed in synonymy
with Lacon.

HISTORY OF THE GENUS. The genus Lacon was established by Castelnau in
1836 for ‘Elater atomarius, varius, fasciatus, etc., de Fabricius [sic].’ Germar
(1840 : 260) redefined the genus and though he credits it to ‘Lap. [Laporte de
Castelnau]’ he does not include the three originally included species. Germar
transferred atomarius to Agrypnus and varius and fasciatus to Adelocera. Germar’s

_ interpretation of Lacon included E. caliginosus Boisduval, E. muticus Herbst,

54 Cs Ma EY von HAVES

L. terrenus Germar, ?E. parviceps Schoenherr, E. crenatus Klug, L. turbidus
Germar, E. murinus Linnaeus, F. vestitus Klug, L. mustellinus Germar, E. ivroratus
Klug, A. crenicollis Ménétriés and also E. ovalis Germar. Thomson (1859 : 103)
designated murinus Linnaeus as the type-species. This interpretation of Lacon was
generally accepted until Hyslop (1921 : 652) drew attention to the fact that murinus
was not originally included in the genus and designated E. atomarius Fabricius
[=Lacon punctatus (Herbst) see p. 76] as the type-species of Lacon.

Hyslop (1921 : 621) remarks that [as a result of his studies] the application of
the names Adelocera and Lacon have been reversed but as he does not make it
clear which species are to be included in each genus, I have accepted Fleutiaux’s
1926 correction of the Schenkling catalogue as the date of publication of the new
combinations.

Golbach (1969) appears to have been unaware of Fleutiaux’s correction to the
Schenkling catalogue as he records as new combinations many of the species
previously transferred by Fleutiaux.

NOTES ON THE GENERA AND SUBGENERA NEWLY PLACED IN SYNONYMY WITH
Lacon. Ocneus Candéze was erected for a single Brazilian species, limbatus
Candéze, in which antennal segments are strongly transverse, the antennal grooves
shallow and the pronotum distinctly bituberculate. Examination of the type-
material (p. 69) has shown that the antennal segments of the male are more strongly
transverse than those of the female. This sexual dimorphism also occurs, though
to a less marked degree, in punctatus Herbst, the type-species of Lacon. The other
diagnostic characteristics of the genus are all of the variable type discussed above.

Scelisus Candéze. Candéze considered this genus to be related to his interpre-
tation of Corymbites. Examination of the type-material of the type-species,
sanguineus Candéze (see p. 81) from Nepal has shown that this species possesses
all the diagnostic characteristics of Lacon. The species differs from L. punctatus
(Herbst) in that the antennal grooves do not extend beyond the mid point of the
sternopleural sutures and the vestiture is setose and not scale-like. These charac-
teristics also occur in L. vitalisi (Fleutiaux) from Laos, and the Madagascan
L. madida (Candéze).

Alaotypus Schwarz was erected for two species from Tonkin (subpectinatus
Schwarz, aspersus Schwarz) in which the antennal grooves are shallow and do
not extend beyond the middle of the sternopleural sutures. The length and depth
of this groove is now known to vary from one species to another and are believed
by the writer to be of no value at the generic level.

Fleutiaux (1918 : 183) placed Alaotypus in synonymy with his interpretation of
Adelocera (Adelocera sensu auctt., nec Latreille) but in his key (1941 : 48) to the
Agrypnitae he treats it as a valid genus. The most recent workers, Ohira
(19690 : 41) and Golbach (1969)-155) also treat Alaotypus as a valid genus, but the
present writer does not consider the diagnostic characteristics to be of sufficient
value to justify the retention of the genus.

Sulcilacon Fleutiaux was established for three species from south-east Asia
(sanguineus (Fleutiaux) = mausoni nom. n., spurcus (Candéze) and geographicus

.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 55

(Candéze)) which possess well defined propleural tarsal grooves. The degree of
development of these grooves is known to vary from one species to another.
Similar grooves are found in the majority of species originally included in
Diphyaulon (q.v.).

Diphyaulon Arnett was proposed as a subgenus of Lepidotus [=Lacon in the
present work] for species in which the pronotum has a pronounced median
longitudinal furrow and ‘usually’ with well defined propleural tarsal grooves.
These characteristics are of value at the specific level only and there is no justifica-
tion for the retention of Diphyaulon, even as a subgenus.

Aulacon Arnett. The three species attributed to this subgenus of Lepidotus
[Lacon of the present work] do not differ from L. punctatus (Herbst) in any
significant way. In my opinion, Arnett’s distinguishing characteristic, the absence
of a pronounced median longitudinal furrow on the pronotum, is of no value at
the generic, or even subgeneric, level.

Zalepia Arnett was erected for those American species with shallow tarsal
grooves on the propleurae and an irregular pronotal surface. These characters
are very variable and of no value at the generic or subgeneric level.

Kobulacon Chujo & Ohira. The type-species guadrinodatus Lewis possesses all
the diagnostic features of Lacon. It differs from the type-species punctatus
(Herbst) only by such very variable characters as the presence of tubercles on
the pronotum and shallow propleural tarsal grooves.

Lepidelater Smith was erected for a single species misticius Mignot. Despite
the author’s comment that the genus resembles Lanelater, the type-species is a
synonym of Lacon subcostata (Candéze).

Arnettia Golbach. The genus is based on an error in the original description
of the type-species. The second antennal segment is small, not large and triangular,
like the third.

Monocyrton Golbach. Only the presence of a posterior median tubercle on
the pronotum distinguishes the members of this subgenus from L. punctatus
(Herbst), the type-species of Lacon. Prothoracic tubercles, either singly or in
pairs, occur in a large number of species and are of no value at the generic or even
subgeneric level.

Cornilacon Golbach. The subgenus is characterized by the antennal grooves
which are deep anteriorly and shallow posteriorly. This character which occurs,
developed to a greater or less degree, in many species throughout the world, is too
variable to justify the retention of the subgenus.

Latilacon Golbach. This South American subgenus was erected on the basis
of the poorly defined propleural tarsal grooves and the presence of tubercles on
the prothorax. The Japanese subgenus Kobulacon was based on the same variable
characteristics.

SEXUAL DIMORPHISM. Members of the genus do not show any marked sexual
dimorphism. In some species, including the type-species, L. punctatus (Herbst),

56 C.-M. EF. von HAYEK

the antennae of the male are a little longer and the segments slightly more transverse
than those of the female.

DISTRIBUTION. The largest number of species occur in the New World and
in the Oriental region and the fewest in Africa. Neboiss (1956) lists a large number
of Australian Lacon species but all those known to me (approximately half Neboiss’s
total) with the exception of modestus (Boisduval) belong to Agrypnus as defined
in this work. JL. modestus (Boisduval) appears to be widespread, though not
common, in the tropics. As far as is known all the other species are comparatively
limited in their distribution.

BIOLOGY AND HABITS. Very little is known of the biology and habits of the
members of this genus.

The adults of the European species are generally found under the bark or
in the decaying stumps of coniferous trees. Adults of modestus (Boisduval)
(Fleutiaux, 1947 : 275) and of certain American species (Kirk, 1922) have been
found under the bark and in wood-borer galleries in the timber of deciduous trees.
The larvae have been found in the same places. Those of L. marmoratus (Fabricius)
are known to feed on Bostrychid larvae (Kirk, 1922) and it seems probable that
the larvae of other species are also predaceous on wood-boring larvae.

SPECIES INCLUDED IN THE GENUS

The following 106 species known to me are congeneric with the type-species
Elater atomarius Fabricius (=Lacon punctatus (Herbst)).

Lacon altaicus (Candéze) comb. rev.

Adeloceva altaica Candéze, 1881 : 2.
Lacon altaicus (Candéze) Fleutiaux, 1926 : 93.
Adelocera altaica Candéze; Tscherepanov, 1957 : 187.

The description is based on an unrecorded number of specimens from Altai
standing as lepidoptera in the Gebler collection. I have not seen this material,
which may be in the MNHN, Paris; ZI, Leningrad (Horn, 1935 : 87); or IRSNB,
Brussels (Fleutiaux, 1945 : 81).

The generic attribution is based on the description.

Lacon antennatus (Lewis)

Adelocera antennata Lewis, 1894 : 27.
Lacon antennatus (Lewis) Fleutiaux, 1926 : 93.

Holotype. JAPAN: 9, Sap. [on underside of card mount = Sapporo]; Adelocera
antennata Lewis, type [Lewis]; Japan. G. Lewis 1910.320 (BMNH).

Lacon argentatus (Candéze) comb. n.
Opatelus argentatus Candéze, 1874 : 42.

LECTOTYPE (present designation). Brazit: ? 9, Bates coll. Ega; Janson
coll. 1903.130; Opatelus argentatus mihi [sic] n. sp. [Janson] Candéze [Gahan]

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 57

(BMNH). The absence of a Candéze determination label may be due to the fact
that Candéze accepted Janson’s name and did not trouble to affix a label himself.

The specimen lacks two features which are characteristic of the genus Opatelus.
The third antennal segment does not resemble the second and there are no tarsal
grooves on the metasternum. It must be assumed that the very small size of
the specimen (4 mm) led Candéze to make this mistake in the generic attribution.

Lacon aspersus (Schwarz) comb. rev.

Alaotypus aspersus Schwarz, 1902 : 309.

Adelocera brevicornis Fleutiaux, 1906b : 211. [Synonymized by Fleutiaux, 1918d : 186.}
Adelocerva adspersa Fleutiaux, 1918d : 186 [unjustified emendation].

Lacon aspersus (Schwarz) Fleutiaux, 1926 : 93.

Alaotypus aspersus Schwarz; Fleutiaux, 1927 : 62.

Alaotypus aspersus Schwarz. Holotype. NortH VIETNAM: ? 9, Tonkin, Montes
Mauson. ? DEI, Eberswalde.

Adelocera brevicornis Fleutiaux. LECTOTYPE (present designation). NortTu
ViETNAM: 9, Tonkin, Montes Mauson, April—Mai, 2300 [metres], H. Fruhstorfer;
Adelocera brevicornis Fleut., Type [Fleut.]; Bull. Soc. Ent. Fr. 1906 p. 211 [Fleut.]
(MNHN, Paris).

This is a distinctive and easily recognizable species and I have therefore accepted
Fleutiaux’s synonymy. Both species are from the same locality and may well be
part of the same series.

Lacon atterimus (Candéze)

Adelocera attevima Candéze, 1889 : 70(4).
Lacon atterimus (Candéze) Fleutiaux, 1926 : 95.

LECTOTYPE (present designation). MADAGASCAR: 3, Madagascar, Museum
Paris; Adelocera atterima Cand. {[Cand.] Candéze vidit (MNHN, Paris).

Lacon auroratus (Say) comb. rev.

Elatey auroratus Say, 1839 : 181.
Lacon auroratus (Say) Fleutiaux, 1926 : 93.
Zalepia aurorata (Say) Arnett, 1969 : II.

LECTOTYPE (present designation). U.S.A.: 9, 931, N.H. [Harris, refers to his
notebook = Elater (Taphiecerus) auroratus S. n.sp./near marmoratus in form of
prothorax] (MCZ, Harvard).

The description is based on an unrecorded number of specimens from New
Hampshire submitted by Dr Harris. There are no specimens in the Say collection
in the ANS, Philadelphia (see p. 279). The specimen measures 13 mm compared
with the published length of ‘eleven-twentieths of an inch’ [= 14.3 mm].

The date of publication of the description is 1839 (not 1836) but it may have
appeared earlier, see note in the list of references p. 295.

I am greatly indebted to Professor Darlington for locating this specimen and
supplying me with the data from Harris’s notebook.

58 C. 1M. EF. von HAYEK

Lacon aurulentus (Candéze)

Adeloceva auvulenta Candéze, 1865 : 6.
Lacon aurulentus (Candéze) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). CEYLON: 9, Janson coll. ex Candeze, 1903.
130; Adelocera aurulenta Cdz. Ceylon, Nietn. [Cand.]; Adelocera aurulenta Cand.
Ceylon [Gahan] (BMNH).

Paralectotype. CEYLON: 1 9, Neitner Ceylon; Janson coll. 1903.130; Adelocera
aurulenta Cdze (El.n.) [Janson] (BMNH).

Lacon avitus (Say) comb. rev.

Elater avitus Say, 1839 : 182.
Lacon avitus (Say) Fleutiaux, 1926 : 93.
Aulacon avitus (Say) Arnett, 1969 : IT.

The description is based on an unrecorded number of specimens from Indiana
[es -A.|

Type-material. As this cannot be found in the ANS, Philadelphia, it must be
regarded as lost. For notes on the Say collection see p. 279.

The generic attribution is based on the following specimen: 1 9, Ky.; avita Say;
Andreas Bolter collection (BMNH).

The description of this species may have appeared before 1839; see note in the
Bibliography, p. 295.

Lacon bicolor Fleutiaux

Lacon (Adeloceva des auteurs) bicoloy Fleutiaux, 1940a : 90.

LECTOTYPE (present designation). Cuina: 3, Yunnan; Lacon bicolor Fleut.,
type. Adelocera [Fleut.] (MNHN, Paris).

Lacon bulwensis (Fleutiaux)

Adeloceva bulwensis Fleutiaux, I9I9 : Io.
Lacon bulwensis (Fleutiaux) Fleutiaux, 1926 : 23.

LECTOTYPE (present designation). TANZANIA: 9, Bulwa, Usumbara; H. Rolle.
Berlin, S.W.11; Adelocera bulwensis Fleut., type [Fleut]; Fleut. Voy. All. et
Jeann. Afr. or. Elat. p. [Fleut.] (MNHN, Paris).

Lacon calabaricus (Candéze)

Adelocera calabarica Candéze, 1874 : 23.
Lacon calabaricus (Candéze) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). NicEr1A: 9, Old Calabar; Janson coll.
1903.130; Ad. calabarica Cdz. n. sp., type [Cand.]; Adelocera calabarica Cand.,
type 2 [Gahan] (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 59

Lacon candezei (Desbrochers des Loges)

Adeloceva candezei Desbrochers des Loges, 1875 : 37.
Lacon candezei (Desbrochers des Loges) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). JORDAN: 3, Naplouse [Nablus]; candezei
m. [Desbr.] (MNHN, Paris).

Schwarz (1894: 145) synonymizes Adelocera kraatzi Schwarz (1893 : 192) from
Jerusalem with candezei. The type-material of kraatzi is probably in the DEI,
Eberswalde (see p. 280). The synonymy has not been confirmed.

Lacon candidus (Fall) comb. rev.

Adelocera candida Fall, 1932 : 59.
Lacon (Zalepia) candidus (Fall) Arnett, 1953 : 7.
Zalepia candida (Fall) Arnett, 1969 : II.

Holotype U.S.A.: 3, Baboquiv. Mts., Ariz. ix/15-30/23; Type. candida [Fall].
M.C.Z. Type 24350 (MCZ, Harvard.)

Lacon carinensis (Candéze)

Adelocera carinensis Candéze, 1891a : 772.
Lacon carinensis (Candéze) Fleutiaux, 1926 : 93.

Holotype. Burma: g, Carin, Ghecu, 1300-1400 m.s.m. Febbraio-Marzo 1888,
legit L. Fea. (MCSN, Genoa according to Binaghi, 1941c : 122. Not confirmed).
Candéze records the locality as: Toungoo; montagnes de Carin, détroit de Ghecu:
1300-1400 metres, mars.

The generic attribution is based on the description.

Lacon castelnaui (Candéze)

Adelocera castelnaui Candéze, 1889 : 71.
Lacon (Monocyrton) castelnaui (Candéze); Golbach, 1969b : 157.

The description is based on two specimens collected at Bahia [Brazil] by Leconte

de Castelnau.

Syntypes: ?IRSNB, Brussels.
The generic attribution is based on the description.

Lacon chabannei (Guérin-Méneville)

Adelocera chabannei Guérin-Méneville, 1829 : pl. 12, fig. 4; 1844 : 41.
Alaus flammula Blanchard, 1841 : pl. 8, fig. 5; 1845 : 136. [Synonymized by Candéze, 1857:
65.]

| Lacon (Monocyrton) chabannei (Guérin-Méneville) ; Golbach, 1969) : 157.

Adelocera chabannei Guérin-Méneville. The material on which Guérin-Méneville
_ based his drawings, which he made under Latreille’s supervision (Cuvier, 1832 : 333)

60 C2. vier SEVANGEAC

has not been located. It may be in the MNHN, Paris or in his own collection in
the IRSNB, Brussels. The published locality is ‘L’interieur du Brazil’.

The generic attribution is based on material in the BMHN which agrees well
with the figure.

Latreille (1834: 144) states that plate 12 was issued in fascicle 4 of Guérin-
Méneville’s work. The appearance of this fascicle is recorded in the Bibliographie
de France, 1829 (November 21) : 783. Fascicles 46-50 containing the text were
published in 1844.

Alaus flammula Blanchard. LECTOTYPE (present designation). BoLivia:
9, A. flammula Blanch. M. D’Orbigney, Chiquitos; type de E. Blanchard [Fleut.]
(MNHN, Paris).

Blanchard records the locality as ‘M. d’Orbigney a pris ce bel insecte sur les
troncs d’arbres dans la province de Guarayos’. Chiquitos is a district in the Llanos
de Guarayos.

The date of publication of this species is generally given as 1837-1843, the date
appearing on the title page of the volume in which the description appeared.
Sherborn and Griffin (1935) who had access to a copy of the whole work in its original
wrappers have shown that volume 6, part 2 appeared in 37 livraisons between
1836 and 1845. The dates on which the figure and description of this species were
published were obtained from Sherborn and Griffin.

Lacon chilensis (Solier)

Agrypnus chilensis Solier, 1851 : 7; pl. 13, fig. I.
Lacon (Aulacon) chilensis (Solier); Golbach, 19696 : 157.

LECTOTYPE (present designation). CHILE: 9, Agrypnus chilensis Sol., Chile
[?Solier]; Type de Solier [Fleut.] (MNHN, Paris).
The published locality is ‘Valdiva cerca de San José.’

Lacon cinctus (Candéze)
Adeloceva cincta Candéze, 1878) : LII (6).

Lacon cinctus (Candéze) Fleutiaux, 1926 : 93.
The description is based on an unrecorded number of specimens from Sumatra.
Type-material: ?>IRSNB, Brussels.

The generic attribution based on the following specimen: 1 9, Marol [?, illegible]
83 : 33 [BMNH registration number = Timor Laut (= Kepuluan Tanimbar)
H. O. Forbes]; Adelocera cincta Candz. (BMNH).

Lacon coeca (Candéze)

Adeloceva coeca Candéze, 1874 : 18.
Lacon (Zalepia) coeca (Candéze); Golbach, 1969b : 159.

LECTOTYPE (present designation). GUATEMALA: 4, Izabal; A. coeca Cdz.
n.sp. type [Cand.]; Guatamala, Sallé coll; Adelocera coeca Cand., Sallé coll. 1413

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 61

[refers to catalogue of Sallé collection where specimen is recorded as the type];
B. C. A. Col. III (1) Adelocera coeca Cand. [Champion] (BMHN).

Lacon cognatus (Candéze)

Adelocera cognata Candéze, 1892a : 796.
Lacon cognatus (Candéze) Fleutiaux, 1926 : 93.

Holotype. ENGGANO: g, Bua-—Bua, collected by Modigliani, May—June, 1891,
and submitted to Candéze by Gestro. (MCSN, Genoa according to Binaghi,
194Ic : 118, Not confirmed.)

Lacon constellatus (Fleutiaux)

Adeloceva constellata Fleutiaux, 1906b : 212.
Lacon constellatus (Fleutiaux) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). JAVA: 9, Java; constellata Fleut., type
[Fleut]; Bull. Soc. ent. Fr. 1906 p. 212 [Fleut.] (MNHN, Paris).

Lacon cribratus (Candéze)

Adelocera cribrata Candéze, 1857 : 63; pl. 1, fig. 7.
Adelocera lacerta Candéze, 1865 : 7. [Synonymized by Candéze, 1874 : 25.]
Lacon cribratus (Candéze) Fleutiaux, 1926 : 93.

Adelocera cribrata Candéze. The description is based on a single specimen
believed to be from Guiana or Brazil in the Guérin-Méneville collection. Eight
years later Candéze (1865 : 7) stated that he believed it to be from Poulo—pinang
[Penang, Malaya], and also (1874: 25) that he has seen a number of specimens
from Malacca in the de Castelnau collection.

Type-material: ?IRSNB, Brussels.

Adelocera lacerta Candéze. LECTOTYPE (present designation). MALAyA: 4d,
Malacca; Janson coll., ex Candéze, 1903.130; Adelocera lacerta Cdz, Malacca Cast.
Cand.]; Adelocera lacerta Cand. Malacca [Gahan] (BMNH).

The species is distinctive and easily recognizable and for this reason I have
accepted Candéze’s synonymy.

Lacon cristatus (Fleutiaux)

Adelocera cristata Fleutiaux, 1918a : 206.
Lacon cristatus (Fleutiaux) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). SouTH VIETNAM: 9, Museum Paris,
Cochinchine, Lemesle, 1865; Adelocera cristata Fleut. co-type [Fleut.] (MNHN,
Paris).

Paralectotype. 9, Sumatra, Palembang; Adelocera cristata Fleut., co-type [Fleut.]
(MNHN, Paris).

62 Cr Ms ES Vion EEA yer

Lacon cuneatus (Candéze) comb. n.
Dilobitarsus cuneatus Candéze, 1865 : 8.

LECTOTYPE (present designation). FRENCH GUIANA: 4, Bar. Cayenne;
Janson coll. 1903 : 130; Dilobitarsus cuneatus Cdze. Type (El.n) (e coll. de
Mniszech) [Janson] (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see
p- 276).

Lacon delagrangei (Buysson)
Adelocera delagranget Buysson, 1891 : 134.
Lacon delagranget (Buysson) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). Syria: 9, Syrie, Amanus C.D. 1891;
Adelocera delagrangei nov. sp. [Buyss.] (MNHN, Paris).

Lacon discoidea (Weber) comb. rev.

Elater discoidea Weber, 1801 : 77.

Elatey pennata Fabricius, 1801 : 239. [Synonymized by Schoenherr, 1817 : 302.]
Lacon discoidea (Weber) Fleutiaux, 1926 : 93.

Zalepia discoidea (Weber) Arnett, 1969 : II.

Elater discoidea Weber. The description was based on an unrecorded number of
specimens from ‘America boreali’. Weber’s types are presumed lost or destroyed
(Lane, in litt.).

Elater pennata Fabricius. LECTOTYPE (present designation). U.S.A.: 9,
[abdomen damaged by Anthrenus] pennata Fab. Caroline, coll. Bosc. [? Bosc, on
green paper] (MNHN, Paris).

Schoenherr regards pennata as the valid name, but since Fabricius (1801 : 252)
refers to Weber under Lucanus piceus, it seems probable that his work appeared
after that of Weber.

The species is distinctive and there is no good reason why the traditional synonymy
should be rejected.

Lacon distinctus (Fleutiaux) comb. rev.

Adeloceva distincta Fleutiaux, 1920a : I14.
Lacon distinctus (Fleutiaux) Fleutiaux, 1926 : 93.
Alaotypus distinctus (Fleutiaux) Ohira, 1970b : 230.
Holotype. Laos: g, Xieng-Khouang, 12.5.19 Vitalis de Salvaza; Adelocera
distinctus Fleut; type [Fleut.] (MNHN, Paris).

Lacon dorsalis (Candéze)

Adeloceva dorsalis Candéze, 1857 : 62; p. I, fig. 4.
Lacon dorsalis (Candéze) Fleutiaux, 1926 : 95.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 63

Syntypes examined. MapaGascar: I ex., Laf. Madagascar; Adelocera dorsalis
Cdze., Cand., type [Janson]; Janson coll. 1903. 130; nov. sp. Madag. Goudot (BMHN).
The absence of Candéze’s determination label is probably due to Janson (see p. 276).
I 2, Collection Chevrolat; dorsalis Cand., type [Fleut.]; Coll. Chevr. co-type Cand.
Mu. [Fleut.]. The specimen stands over a blue Chevrolat label: Adelocera dorsalis
Cand., Mon. 1, 1857, p. 42. 16. Madagascar (MNHN, Paris). There is no Candéze
determination label. It may have been removed by Chevrolat.

Candéze also refers to material in the Mniszech collection. ? IRSNB, Brussels.

Fleutiaux (1903) : 263) described var. obscuripennis based on an unrecorded
number of specimens from: Region des Diego Suarez, montagne d’Ambre. This
material has not been located. ? MNHN, Paris.

According to Fleutiaux (1907a : 162), Adelocera auricollis Schwarz (19050 : 273)
described from the same locality as var. obscuripennis is a synonym. Syntype-
material: ? DEI, Eberswalde. The synonymy has not been confirmed.

Lacon drusa (Marseul) sp. rev.

Adeloceva drusa Marseul, 1870 : 380.
Adelocera bruleriei Desbrocheres des Loges, 1875 : 37. Syn. n.

Adelocera drusa Marseul. LECTOTYPE (present designation). Syria: 4,
Adelocera drusa Kar-Eliz [sic] remainder of label illegible [Marseul]; Museum Paris.
coll. de Marseul, 1890 (MNHN, Paris). The published locality is Kab-Elias, Syria.

Adelocera bruleriei Desbrocheres des Loges. LECTOTYPE (present designation).
LEBANON: @, Liban; Brulerie [sic] m. [Desbrocheres] (MNHN, Paris).

Adelocera drusa Marseul is not a synonym of A. graeca Candéze. Schenkling’s
(1925 : 10) synonymy is presumably based on Candéze’s (1874 : 21) comment that
drusa resembles graeca.

Lacon dubius Candéze comb. rev.

: Lacon dubius Candéze, 1857 : 160; pl. 2, fig. 6.
| Pyrganus dubius (Candéze) Golbach: 1968: 198.

- LECTOTYPE (present designation). CoLomsBiA: 9, Columbia [sic., Janson];
: Janson coll. 1903. 130; Lacon dubius Cand. e coll. Lacord. Candéze, type [Janson];
L. dubius Cand. type [Cand.] (BMNH).

The published locality is ‘Colombie’ [S. America]. The discrepancy between
| the published locality and the label is probably due to Janson.

The structure of the fourth tarsal segments, which are strongly oblique at the
apex, suggests that the retention of Lacon and Dilobitarsus as separate genera is
not entirely justified. However, this problem cannot be resolved until more material
becomes available.

Lacon expansus (Fleutiaux)

| Adeloceva expansa Fleutiaux, 1920a : 114.
| Lacon expansus (Fleutiaux) Fleutiaux, 1926 : 93.

|

|

64 Cl 2M. BS von HAYEK

Holotype. N. VietTNAM: 9, Hoa Binh, Tonkin Vitalis, Janv. 17; Hoa Binh,
Tonkin de Cooman; non Vitalis sed P. de Cooman, Adelocera expansa Fleut., type
[Fleut.] (MNHN, Paris).

Lacon fleutiauxi (Schwarz)

Adelocera fieutiauxt Schwarz, 1902a : 311.
Lacon fleutiauxi (Schwarz) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). EQUATORIAL GUINEA: 3, Benito, Congo
Franc.; Adelocera fleutiauxi n. sp. [Schw.]; fleutiauxi Schwarz, type [Fleut.];
Sulcilacon fleutiauxi Schw., type ¢ [Girard] (NMHN, Paris).

The transference of this species to Sulcilacon has not been published.

Lacon foveatus (Candéze)

Adelocera foveata Candéze, 1895) : 52.
Lacon foveatus (Candéze) Fleutiaux, 1926.93.

The description is based on a single specimen from Tamatave, Madagascar,
collected by Alluaud between April and August, 1893.

Holotype: The specimen cannot be found in the MNHN, Paris. ? IRSNB,
Brussels.

The generic attribution is based on a note under olsoufieffi Fleutiaux in the
Fleutiaux collection (MNHN, Paris) ‘foveata Cand. comparé au type, Janv. ’42.’.
The synonymy has not been published or confirmed.

Lacon fulvipennis Fleutiaux
Lacon fulvipennis Fleutiaux, 1932a : 49.

Holotype g. Mapacascar: Museum Paris, Madagascar, Prov. de Fénérive, Reg.
de Soaniérana, A. Mathieux 1905; fulvipennis Fleut., type [Fleut.] (MNHN, Paris).

Lacon funebris (Solsky)

Adelocera funebris Solsky, 1881 : 231.
Lacon funebris (Solsky) Fleutiaux, 1926 : 3.

The description is based on an unrecorded number of specimens from ‘valle
Sarafschan’ [= Zeravshan, TADZHIKSTAN S.S.R].

The type-material is probably in the ZMV, Moscow (E. B. Britton in litt.).

The generic attribution is based on the following specimen: TURKESTAN,
Q-15.vii.1960, determined by Skopin (BMNH).

Schenkling (1925: 9) records the date of publication of this species as 1882,
apparently referring to a separate with pagination 1 —. The separate is unknown
to me. There seems to be no reason why the date on the title-page of the Hovae
Soc. ent. ross. '? should not be accepted.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 65

Candéze (1891 : 13) records Adelocera tristis Kraatz (1882 : 319) as a synonym
of funebris. The synonymy has not been confirmed.

Adelocera tristis Kraatz. The description is based on an unrecorded number of
specimens (‘ziemlich selten’) collected by Haberhauser near Samarkand.
Type-material: ?Kraatz collection, DEI, Eberswalde (Horn, 1935; 141).

Lacon ganglbauri (Schwarz)

Adelocera ganglbauri Schwarz, 1894 : 145.
Lacon ganglbauri (Schwarz) Fleutiaux, 1926 : 93.

The description is based on two specimens from Beirut [LEBANON], Appl., 1878
in the NM, Vienna.

The generic attribution is based on the description.

Lacon geographicus (Candéze) comb. rev.

Adelocera geographica Candéze, 1865 : 7.
Lacon geographicus (Candéze) Fleutiaux, 1926 : 93.
Sulcilacon geographicus (Candéze) Fleutiaux, 1941¢ : 48.

LECTOTYPE (present designation). BornrEo: 3g, Borneo 1426; Sar. 874;
Janson coll. 1903.130; Adelocera geographica Cdz. [Cand.]; Adelocera geographica
Cand., type [Gahan] (BMNH).

Lacon graeca (Candéze)

Adeloceva graeca Candéze, 1857 : 61.
Lacon graeca (Candéze) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). 4g, graeca type [Fleut.]; graeca Cand.
type, Collection Chevrolat; [Fleut.]._ The specimen stands beside a Chevrolat label:
Adelocera graeca (Chev.) Cand. Mon.1. 1857 p. 61 13. Coll. Chevrolat (MNHN,
Paris.)

The published locality is ‘Gréce’ [GREECE].

Lacon impressicollis (Say) comb. rev.

Elatey impressicollis Say, 1825 : 260.

Adelocera senilis Germar, 1840 : 259. [Synonymized by Candeze, 1859 : 58.]
Adelocera impressicollis (Say) LeConte, 1853 : 490.

Lacon impressicollis (Say) Fleutiaux, 1926 : 490.

Zalepia impressicollis (Say) Arnett, 1969 : IT.

Elater impressicollis Say. Say proposed this name for Elater fuscus Melsheimer
(1806 : 44, nomen nudum) which he regarded as preoccupied, presumably by
Elater fuscus Fabricius (1801). Say does not record a locality or state whether he
based his description on Melsheimer’s specimens or on material in his own collection.
Say’s own material is presumed lost (see p. 279). Horn (1936: 127) states that

E

66 CoM ES Vion EDAYCR ES

Melsheimer’s collection is in the MCZ Harvard but up to the present time the
specimens have not been located.

The interpretation of the species is based on a specimen determined by LeConte:
I g, yellow disk (= Western States); A. impressicollis (Say) LeC. lepturus Say,
senilis Germar [LeConte]; Adelocera impressicollis Say (MCZ, Harvard).

Adelocera senilis Germar. LECTOTYPE (present designation). U.S.A.: g, H
fon green triangular label, ? = Hertz, see below]; North America; Janson coll.
ex Schaum, 1903.130; senilis m. lepturus Say. Amer. bor. 879? [Germar] (BMNH).

Germar received the material from Virginia, on which he based his description,
from Zimmerman and Hertz. The appearance of the name ‘lepturus Say’ on the
label is puzzling. The explanation may be that Germar learned of LeConte’s
(1859a ; 620) synonymy and altered or replaced his original determination label.

LeConte (1859a@ : 182) states that E. lepturus Say (1836: 128) does not differ
from wmpressicollis. The description of Elater lepturus is based on an unrecorded
number of specimens from ‘Pennsylvania, Indiana and North Carolina. Harris’.
As the material cannot be found in the Say collection (see p. 279) in the ANS,
Philadelphia or the Harris collection (see p. 276) in the MCZ, Harvard, the type
material must be presumed to be lost. The synonymy cannot therefore be con-
firmed.

Lacon inaequalis (Candéze)

Adeloceva inaequalis Candéze, 1857 : 67.
Lacon (Aulacon) inaequalis (Candéze); Golbach, 19696 : 157.

LECTOTYPE (present designation). CoLomBiaA: 9, Columb. [sic] Dup. 29;
Collection Chevrolat; inaequalis Cand., type [Fleut.]; Adelocera inaequalis (Chev.)
Cand., type Mon.1. 1857 p. 67. 21 Nova Grenada. D. Dupont [Chevrolat] (MNHN,
Paris).

The published locality is ‘Nouvelle Grenade’ the name by which Colombia was
known between 1831 and 1861.

Lacon incommodus Fleutiaux
Lacon incommodus Fleutiaux, 1934é : 364.

LECTOTYPE (present designation). PHILIPPINES: 3, Davao, Mindanao, Baker;
Lacon incommodus Fleut., type ex Adelocera [Fleut.] (MNHN, Paris).

This species has never been included in the genus A delocera.

Lacon incomptus (Kraatz)

Adeloceva incompta Kraatz, 1832 : 319.
Lacon incomptus (Kraatz) Fleutiaux, 1926 : 93.

The description is based on an unrecorded number of specimens collected by
Haberhauser near Samarkand [UZBEck S.S.R.].

Type-material: ? Kraatz collection, DEI, Eberswalde (Horn, 1935 : 141).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 67

The generic attribution is based on Kraatz’s comment that the species bears a
very close resemblance to tvistis Kraatz (see funebris Solsky, p. 65).

Lacon inflatus (Candéze)

Adelocera inflata Candéze, 1857 : 62.
Lacon inflatus (Candéze) Fleutiaux, 1926 : 95.

LECTOTYPE (present designation). MapAcascar: 9, Deyr. Madagascar; Ad.
inflata [Cand.]; Adelocera inflata Cdze, Cand., type (e coll. Deyrolle) [Janson]
(BMNH).

Paralectotypes: 2 ex., Laf. Madagascar; Adelocera inflata Cdze, Cand. [Janson];
A. nigromicans Gory in mus. (e coll. de Laferté) [Janson]. One specimen bears an
additional label = nigromicans Gory Madagas. [Gory] (BMNH).

Lacon jacquieri (Candéze)

Adeloceva jacquieri Candéze, 1857 : 64; p. I. fig. 8.
Lacon (Monocyrton) jacquieri (Candéze); Golbach, 1969b : 15.

LECTOTYPE (present designation). FRENCH GUIANA: 9, Dej. Cayenne;
Adelocera jacquieri (Dej.) Cdze, Cand., type [Janson]; Agryp. jacquieri Dej. Cat.
(e coll. Dejean) [Janson] (BMNH).

Paralectotype: 9, labels as lectotype, but without the word ‘type’ on Janson’s
label.
The absence of Candéze’s determination label is probably due to Janson (see p. 276).

Lacon jeanvoinei Fleutiaux
Lacon jeanvoinei Fleutiaux, 1941b : 184.
LECTOTYPE (present designation). N. VieEtTNAM: 9, Chapa, Tonkin, ex
Jeanvoine coll. Clermont; Lacon (Adelocera) jeanvoinei Fleut., type [Fleut.] (MNHN,
Paris).

Paralectotype. 9, labels as lectotype but without Fleutiaux’s determination
label (MNHN, Paris).

Lacon laconoides (Schwarz) comb. n.
Scelisus laconoides Schwarz, 1902a : 343.

The description is based on two specimens from N. VIETNAM: Tonkin, Montes
Mauson.

Syntypes: ? DEI, Eberswalde.

The generic attribution is based on the description and the figure in Schwarz
(1907 : pl. 6, fig. 7).

68 Ci Me FS von HAW ETS

Lacon laoticus Fleutiaux
Lacon laoticus Fleutiaux, 1927 : 68.

LECTOTYPE (present designation). Laos: 3, Ht. Mekong, Ba ni Hona 25.3.20;
Lacon laoticus Fleut., type [Fleut.] (MNHN, Paris).

Lacon laticollis (Candéze)

Elater pulverulentus Fabricius, 1801 : 228. [Junior primary homonym of Elater pulverulentus
Panzer, 1795.|

Adelocera laticollis Candéze, 1857: 59. Syn.n.

Adelocera abervans Candéze, 1874 : 23. Syn. n.

Aynettia abervvans (Candéze) Golbach, 1969b : 155.

Lacon (Latilacon) laticollis (Candéze); Golbach, 19696 : 158.

Elater pulverulentus Fabricius. LECTOTYPE (present designation). GUYANA:
dg, small green label identifying Fabrician specimen (Zimsen, 1964: 12) Type
[curatorial label]; Essequibo. Smidt. Mus. D. Sehestedt. Elater pulverulentus Fab.
[mss. label replacing Fabricius’ label, see Zimsen, loc. cit.] (ZMU, Copenhagen).

The published locality is ‘America meridionali. D. Smidt. Mus. D. de Sehestedt’.

This species has not appeared in any catalogue or other work since Schoenherr
(1817 : 280).

Adelocera laticollis Candéze. Holotype. FRENCH GuIANA: 4, Laf. Cayenne;
Janson coll. 1903 : 130; Adelocera laticollis Cdze, Cand., Type (e coll. de Laferté)
[Janson]; nov. sp. Cayenne, Mus. Reiche (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see p. 276).

Lacon laticollis (Candéze, 1857 : 59) is a junior secondary homonym of Lacon
laticollis Candéze, 1857: 146 = Agrypnus laticollis (Candéze) (see p. 274) but as
the two species are not congeneric the junior name does not require replacement
(IRZN, Art. 59 (b)).

Adelocera aberrans Candéze. LECTOTYPE (present designation). VENEZUELA:
Q, Caracas; A. aberrans Cdz. type [Cand.]; Ex Musen A. Sallé 1897; Museum Paris,
1952. Coll. R. Oberthur (NMHN, Paris).

Candéze’s statement concerning the size of the second antennal segment is
erroneous. It is not large and triangular like the third, but very small and
moniliform. As the type material until recently stood unrecognised in the MNHN,
Paris, it seems probable that Golbach based his interpretation of the species, for
which he established the genus Arnettia, on Candéze’s misleading description and
not on authentically identified material.

Lacon lepidopterus (Panzer) comb. rev.

Elater lepidoptera Panzer, 1801, part 76, no. 4.
Lacon lepidoptera (Panzer) Fleutiaux, 1926 : 93.
Adelocera lepidopteva (Panzer); Tscherepanov, 1957 : 189.

The description is based upon material ‘Lectus semel Berolini [Berlin. EAST

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 69

GERMANY] in muro, mens. Jun. 1798 a Dom. Funk Pharmaop. Gsresensi—Baruth,
qui eundem benevole mecum communicavit.’ I have been unable to trace this
material.

The generic attribution is based on material standing over this name in the
major European collections.

Lacon limbatus (Candéze) comb. n.
Ocneus limbatus Candéze, 1857 : 85; pl. 2, fig. 27.

LECTOTYPE (present designation). Brazit: g, Laf. Brazil; Janson coll.
1903 : 130; Ocneus limbatus [Cand.]; Ocneus limbatus Cdz. Cand. type ¢$ [Janson]
(BMNH).

Paralectotype. dg, labels as lectotype but without Candéze’s determination
label (BMNH).

The lectotype, paralectotype and the seven other specimens known to me all
differ from Candéze’s generic diagnosis in that the posterior coxal plates are not
wider laterally than in the mid line.

Lacon linearis (Candéze)

Adelocera linearis Candéze, 1865 : 6.
Lacon (Aulacon) linearis (Candéze); Golbach, 19696 : 93.

LECTOTYPE (present designation). FRENCH GUIANA: 3, Janson coll. 1903.130;

Janson coll. ex Candéze; Adelocera linearis Cdz., Cay. De B. [Cand.]; Adelocera
linearis Cand., Cayenne [Gahan] (BMNH).

Lacon lithophilus (Candéze)

Adeloceva lithophila Candéze, 1857 : 60.
Lacon lithophilus (Candéze) Fleutiaux, 1926 : 94.

The description is based on an unrecorded number of specimens from EGYPT
received with this name from Dohrn. The BMNH collection does not possess
any specimens with Dohrn’s determination label or known to come from his
collection.

Type-material: ?IRSNB, Brussels (Candéze collection) or IZPAN, Warsaw
(Dohrn collection, see p. 274).

The generic attribution based on the following specimen: Adelocera lithophila
Cdz. Egypt. Schaum [Cand.]; Janson coll. ex Candéze, 1903.130 (BMNH).

Lacon longicornis (Champion)

Adeloceva longicornis Champion, 1894 : 261.
Lacon (Cornilacon) longicornis (Champion); Golbach, 1969) : 158.

LECTOTYPE (present designation). GUATEMALA: 4, Capetillo, Guatemala;
B.C.A. Col. 111(1) Adelocera longicornis Ch. g¢ [Champ.]; sp. figured (BMNH).

70 CC. M. FE. von HAYEK

Paralectotypes: 4 9, with same locality and determination labels as the lectotype
(BMNH).

Lacon luzonicus (Candéze)

Adelocerva luzonica Candéze, 1875 : CXIX.
Lacon luzonicus (Candeze) Fleutiaux, 1926 : 93

The description is based on 1 g and 1 2 from Luzon.
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on Candéze’s comment that the species resembles
A. aurulenta Candéze (= Lacon, see p. 58).

Lacon maculatus (LeConte) comb. rev.

Adeloceva maculata LeConte, 1866 : 389.
Lacon maculatus (LeConte) Fleutiaux, 1926 : 93.
Zalepia maculatus (LeConte) Arnett, 1969 : II.

LECTOTYPE (present designation). U.S.A.; 9, Pen.; Type 2376 [MCZ curatorial
label]; A. maculata Lec. [LeC.] (MCZ, Harvard).

Paralectotype: 9, D.C.; Adelocera maculata LeC. [Ulke] (CM, Pittsburgh, Ulke
collection).

The description is based on two specimens, one from Philadelphia [Pennsylvania]
found by J. Johnson Brown and another from Washington, D.C., by Mr Ulke.
Ulke appears to have been in the habit of affixing his own determination labels on
specimens described by LeConte. See also Lacon pyrsolepis LeConte p. 78.

Lacon madidus (Candéze)

Adelocera madida Candéze, 1889 : 70 (4).
Lacon madidus (Candéze) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). MADAGASCAR: 9 Museum Paris, Mada-

gascar, Humblot 1885; 6364, 85; Adelocera madida Cand. sp. n [Cand.] (MNHN,
Paris).

Lacon maklini (Candéze) comb. rev.

Adeloceva maklini Candéze, 1865 : 6.
Lacon maklimi (Candéze) Fleutiaux, 1926 : 93.
Adelocera (Compsolacon) maklini (Candéze); Van Zwaluwenburg, 1966 : 298.

The description is based on an unrecorded number of specimens from Japan
received from Professor Maklin of Helsingfors University. Van Zwaluwenburg
(1957 : 10) records a specimen ‘which is probably the type’ in the ZMU Helsinki.

The generic attribution is based on the following specimen: 2, Japan G. Lewis,
1910 : 320; Adelocera maklini Cdze [Janson] (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 7p

Lacon mamillatus (Candéze)

Adelocera mamillata, Candéze, 1865 : 7.
Lacon (Aulacon) mamillata (Candéze); Golbach, 1969b : 157.

LECTOTYPE (present designation). FRENCH GUIANA: Q, Bar. Cayenne,
Adelocera mamillata Cdze, type (El. n.) (e coll. de Mniszech) (BMNH). The absence
of Candéze’s determination label is probably due to Janson (see p. 276).

Lacon marmoratus (Fabricius) comb. rev.

Elater marmorata Fabricius, 1801 : 227.
Lacon marmorata (Fabricius) Fleutiaux, 1926 : 95.
Diphyaulon marmorata (Fabricius) Arnett, 1969 : II.

LECTOTYPE (present designation). U.S.A.: 9, marmorata Fab. Caroline,
Coll. Bosc [?Bosc on green paper] (NMHN, Paris).

Fabricius records the location of the material as Mus. D. Bosc. Zimsen (1964 : 158
No. 2629) records a specimen in the Kiel collection. This may be a syntype retained
by Fabricius, or a subsequent addition to the collection (not examined).

Lacon mausoni nom. n.

Adelocera sanguinea Fleutiaux, 1908 : 164.

Sulcilacon sanguineus (Fleutiaux) Fleutiaux, 1941c : 48.

Lacon sanguineus (Fleutiaux) comb. n. [Secondary junior homonym of Lacon sanguineus
(Candéze, 1863.)]

LECTOTYPE (present designation) N. VietNam: 9, Tonkin, Monts Mauson,
Avril-Mai, 2—3000 H. Fruhstorfer; Adelocera sanguinea Fleut., type [Fleut] Bull
Soc. Ent. Fr. 1908, p. 164 [Fleut.] (MNHN, Paris).

Lacon mexicanus (Candéze)

Adelocera mexicana Candéze, 1857 : 70.
Aulacon mexicanus (Candéze) Arnett, 1969 (March) : 11.
Lacon (Aulacon) mexicana (Candéze); Golbach, 1969b (November) : 157.

Syntype examined. Mexico: g, Mexique Gehin; Collection Chevrolat; Mexicana
Cand., type [Fleut.]; Adelocera mexicana Chevr. Cand. Mon. I. p. 70, 25, 1857
Mexico, D. Gehin, Coll. Chevrolat [Chevrolat] (MNHN, Paris).

Van Dyke (1943 : 44) records that he examined Candéze’s type in the IRSNB,
Brussels in 1932. This is presumably Candéze’s second specimen which was
submitted by Mniszech (not examined). Van Dyke was mistaken in his belief
that nobilis Candéze (see p. 72) is a synonym of mexicanus.

Lacon modestus (Boisduval)
Elatey modestus Boisduval, 1835 : 108.

Agrypnus nigroplagiatus Blanchard, 1853: 85; pl. 6, fig. 7. [Synonymized by Candéze,
1874 : 27.]

72 CoM. ES VoNeE AY EEK

Lacon modestus (Boisduval) Fleutiaux, 1926 : 94.

Lacon coomani Fleutiaux, 1927 : 66.

Lacon modestus var. coomani Fleutiaux; Fleutiaux, 1947 : 275.
Zalepia modesta (Boisduval) Arnett, 1969 (May) : 11.

Lacon (Zalepia) modesta (Boisduval); Golbach, 1969 (November) : 159.

Elater modestus Boisduval. Lectotype (Van Zwaluwenburg, 1959 : 355).
AUSTRALIA: 9, Dej. N. Holl.; Agrypn. modestus Boisd. Dej. Cat. (e coll. Dejean)
[Janson]; Adelocera modesta Boisd. Cand., type. Cdze [Janson] (BMNH).

Paralectotypes: 2 9, labels as lectotype but without the word ‘type’ on the
determination labels (BMNH).

Boisduval based his description on an unrecorded number of specimens from
‘Nouvelle Hollande’ [Australia] in the Dejean collection (see p. 274). The absence
of Boisduval’s and/or Dejean’s labels is probably due to Janson (see p. 276).

Agrypnus mnigroplagiatus Blanchard. LECTOTYPE (present designation).
Borneo: 9, Borneo, Bandjermassin, Jacquinot 1841: Museum Paris, Agrypnus
nigroplagiatus [? Blanchard]; type de Blanchard [Fleut.]; Adelocera modesta
Boisd. (MNHN, Paris).

Lacon coomani Fleutiaux. LECTOTYPE (present designation). N. VIETNAM: 9,
Hoa Binh. A. de Cooman; Lacon coomani Fleut., type [Fleut.] (MNHN, Paris).

Fleutiaux (1947 : 275) gives a useful review of the synonymy and biology of this
species.

The description, distribution and size (5 [presumably German] lines, 12 mm) of
Agrypnus cribrosus Eschscholtz (1829 : 32) suggests that this species may be
conspecific with Lacon modestus (Boisduval), the only member of the subfamily
Agrypninae recorded from Hawaii (Sharp & Scott, 1908 : 368).

Eschscholtz based the description of cribrosus on an unrecorded number of
specimens from ‘Sandwichii insulae’ [Hawaii]. The type-material has not been
located (for history of the Eschscholtz collection see p. 274). Candéze (1857 : 161)
comments that cribrosus resembles Agrypnus caliginosus Guerin but the distribution
of this species appears to be restricted to Australia. Agrypnus cribrosus Eschscholtz
is not recorded in the Schenkling (1925) catalogue, or Fleutiaux’s (1926) corrections
thereto.

Lacon monticola (Candéze)

Adelocera monticola Candéze, 1897 : 6.
Lacon monticola (Candéze) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). ¢, Himal. Coll. Jekel; monticola Cand.,
n.sp [Cand.]; monticola Cand., type. El. nouv. VI, p. 6 [Fleut.] (MNHN, Paris).

The published locality is ‘Himalaya’ without further details.

Lacon nobilis (Fall) comb. n.

Adelocera nobilis Fall, 1932 : 58.
Aulacon nobilis (Fall) Arnett, 1969 : 11

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 73

LECTOTYPE (present designation). U.S.A.: 9, Baboquiv. Mts. Ariz. 7/1-15/24;
N.B.XII p. 137; Type nobilis [? Fall]; (MCZ, Harvard). Length: 20-5 mm.

Paralectotypes: 2 9, same locality and date as lectotype. I 3, same locality,
dated: 8/1-15/24 (MCZ, Harvard). None of the specimens bear determination
labels.

Comparison of the lectotype of nobilis with the syntype of mexicana (Candéze)
(see p. 71). has shown that Arnett’s (1952 : 114) statement that these two species
are distinct is fully justified.

Lacon novus Fleutiaux
Lacon novus Fleutiaux, 1934e : 364.

LECTOTYPE (present designation). PHILIPPINES: 3g, Mt. Banahao, Luzon;
Lacon novus Fleut., type ex Adelocera [Fleut.] (MNHN, Paris).

Lacon obscurus Fleutiaux
Lacon (Adeloceva des Auteurs) obscurus Fleutiaux, 19404a : 90.

LECTOTYPE (present designation). Curva: g, Yunnan Sen, 1911; Lacon
(Adelocera) obscurus Fleut., type [Fleut.] (MNHN, Paris).

Lacon oliveri (Candéze)

Adelocera oliveri Candéze, 1874 : 20.
Lacon olivieri Schwarz, 1906 : 11 [Unjustified emendation.]
Lacon oliveri (Candéze) Fleutiaux, 1926 : 93.

Holotype. Inp1a: 9, Darjeeling, India; Janson coll. 1903.130; Adelocera oliveri
(Janson), type [ Janson] ms., Candéze [Gahan] (BMNH).

Lacon olsoufieffi Fleutiaux
Lacon olsoufiefi Fleutiaux, 1932f : 450.

Holotype. g, Mapacascar: Madagascar, Perinet; Lacon olsoufieffi Fleut., type
[Fleut.] (MNHN, Paris).

The specimen also bears a note in Fleutiaux’s handwriting ‘foveata Cand.
comparé au type, Janv. 42’. The synonymy has not been published or confirmed.

Lacon orientalis (Fleutiaux)

Adelocera orientalis Fleutiaux, 1918d : 185.
Lacon orientalis (Fleutiaux) Fleutiaux, 1926.

LECTOTYPE (present designation). NorTH VIETNAM: 3, Chapa, Tonkin,
Vitalis 28.5.16; Adelocera orientalis Fleut. Type [Fleut.] (MNHN, Paris).

74 ClaMi Be vione AWE KK

Paralectotype: g, same locality as lectotype but dated Mai 1916 and without
an identification label (MNHN, Paris).

Lacon palliatus (Latreille)
Elatey palliatus Latreille, 1809 : 230; pl. 16, fig. 3.

Adeloceva chapuisiit Candéze, 1857 : 66; pl. 1, fig. 5. [Synonymized by Candéze, 1874 : 29.]
Lacon (Monocyrion) palliata (Latreille); Golbach, 19696 : 157.

The date of publication of Elater palliatus Latreille is usually recorded as
1811: 145. This is the date on the title page of the re-issue of Humboldt and
Bonpland’s Voyage ... etc. This work, and especially the 1809 edition, is
very scarce and was probably unknown to many workers including Candéze
(see below).

The present location of the material collected by Humboldt and Bonpland is
unknown to me. I consider it reasonable to assume that Latreille retained at
least one specimen of the new species he described and that this specimen is now
in the BMNH collection (see below).

Elater palliatus Latreillee LECTOTYPE (present designation). MExIco: J,
Dej. Mexico; Janson coll. 1903.130; Ad. chapuisii Cdze, Cand., Type; Agrypn.
palliatus Dej. Cat. (e coll. Dejean) e mus. Latreille; Adelocera palliata Latr. (Elater
palliatus Latr.) [locality and all determination labels in Janson’s handwriting]
(BMNH).

The published locality is “Trouvé parmi des gramineés pres de volcan de Jorillo,
dans le Nouvelle Espagne’ [Mexico]. The specimen agrees well with the figure,
especially as regards the markings on the elytra. The absence of Latreille’s
determination label is probably due to Janson (see p. 276).

Janson’s labels indicate that he believed this specimen to be the type of 4.
chapuisit Candéze. This cannot be the case as the colour pattern on the elytra does
not agree with Candéze’s figure.

Adelocera chapuisu Candéze. The description is based on an unrecorded number
of specimens from Mexico submitted by de la Ferté Sénectére.

Type-material. There are no specimens in the BMNH determined by Candéze
or from the de la Ferté Sénectére collections which agree with Candéze’s figure.
?IRSNB, Brussels (see p. 271).

Candéze lists Agrypnus palliatus De}. ed. 3. p. 99 as a Synonym of chapuisu,
though Dejean (1833: 99) credits the species to Latreille and almost certainly had the
type-material in his collection (see above and p. 277, Latreille collection). Candéze
probably did not know of Latreille’s work in which palliatus was described. I believe
that he based his synonymy on the following specimen: 3, Dej. Columbia [sic,
Janson]; Adelocera palliata Latr. (Elater palliatus Latr); A. chappuisii Cdze,
Agrypnus palliatus Dej. cat. (e coll Dejean) [both determination labels by Janson]
in the BMNH. The markings of this specimen agree well with Candéze’s figure.
The markings of the elytra vary considerably within the species and I believe that
the synonymy is fully justified.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 75

Lacon parallelus (Lewis)

Adelocera parallela Lewis, 1894 : 28.
Lacon parallelus (Lewis) Fleutiaux, 1926 : 93.

LECTOTYPE (Present designation). JAPAN: 9, Sap. [underside of card mount];
Sapporo; Japan, G. Lewis, 1910: 320; Adelocera parallela Lewis, type [Lewis]
(BMNH).

Lacon pectinatus (Candéze)

Adelocera pectinata Candéze, 1865 : 8.
Lacon (Cornilacon) pectinata (Candéze); Golbach, 19696 : 158.

Holotype. FRENCH GUIANA: J, Bar. Cayenne, Janson Coll. 1903 : 130; Adelocera
pectinata Cdze, type (El. n.) (e coll. de Mniszech) [Janson] (BMNH).
The absence of Candéze’s determination label is probably due to Janson (see p. 276).

Lacon pectinicornis (Champion)

Adeloceva pectinicornis Champion, 1894 : 262; pl. To, figs 5, 5a, 5b.
Lacon (Cornilacon) pectinicornis (Champion); Golbach, 1969b : 158.

Holotype. Nicaracua: g, Chontales, Nicaragua, T. Belt.; B.C.A. Col. III (1)
Adelocera pectinicornis Ch. 3 (Champ.); sp. figured (BMNH).

Lacon pectoralis (Fairmaire)

Adelocera pectovalis Fairmaire, 1884b : 276.

Lacon pectoralis (Fairmaire) Fleutiaux, 1926 : 93.
The description is based on an unrecorded number of specimens from MADAGASCAR.
Type-material: not found in MNHN, Paris. ? IRSNB, Brussels (see p. 271).

The generic attribution based on Fairmaire’s statement that the species
resembles inflata Candéze (p. 67).

Lacon philippinus (Fleutiaux)
Lacon (Adelocera) philippinus Fleutiaux, 1934c : 478.

Holotype. PuitipPinEs: 9, Kabasalan, Zamboanga, Mindanao, P.1.39.32; H. C.
Muzzal Collector, F. C. Haddon collection; Lacon (Adelocera) philippinus Fleut.,
Type [Fleut.] (MNHN, Paris).

Fleutiaux’s use of (Adelocera) in this context is intended not as the name of a
subgenus but to indicate that his interpretation of Lacon corresponds to Adelocera
of other authors. See also Fleutiaux (1940a).

Lacon pictus (Fleutiaux)

Adelocera picta Fleutiaux, 1902 (April) : 112.
Lacon pictus (Fleutiaux) Fleutiaux, 1926 : 93.
Lacon (Monocyrton) picta (Fleutiaux); Golbach, 1969b : 157.

76 COMES ES Vion at A

LECTOTYPE (present designation). Brazit: g, Jatahy, Prov. Goyas, Brasil.
Sept. a Nov. 97; Adelocera picta Fleut., type [Fleut.]; Bull. Soc. ent. Fr. 1902
p. 112 [Fleut.] (MNHN, Paris). Length: 13 mm.

Paralectotypes. 2 g, same locality as lectototype. Length: 13-6 mm and
12mm. 19,1, Jatahy, Goyas, Brasil (MNHN, Paris). Length: 3, 9:5 mm, 2
16-8 mm.

Fleutiaux (1907a : 162) synonymizes Anacantha bicostata Schwarz (1902) (July) :
196) with this species. The synonymy has not been confirmed. Schwarz’s descrip-
tion is based on an unrecorded number of specimens from Brazil, Prov. Goyas,
Jatahy. Syntypes: ? DEI, Eberswalde.

Lacon pollinaria (Candéze)

Adelocera pollinaria Candéze, 1857 : 68; pl. I, fig. 14.
Adelocera adspersa Candéze, 1874 : 30. [Synonymized by Champion, 1894 : 260.]
Lacon (Aulacon) pellinaria (Candéze); Golbach, 19696 : 157. [Unjustified emendation.]

Adelocera pollinaria Candéze. LECTOTYPE (present designation). BRaziL:
2, Dej. Brasilia; Janson coll. 1903 : 130; Adelocera pollinaria (Dej.) Cdze, Cand.,
type (Janson); Agrypnus pollinarius Dej.Cat. (e coll. Dejean) [ Janson] (BMNH).

Adelocera adspersa Candéze. Holotype. GUATEMALA: 4g, Guatemala, Salle
coll.; Isabal; Adelocera adspersa Cand. Salle coll. 1414; A. aspersa [sic] Cdz, type
[Cand]; pollinaria Cand. [Champ.]; sp. figured; B.C.A. Col. III (1) Adelocera pollinaria
Cand. [Champ.] (BMNH).

Lacon punctatus (Herbst)

Elater punctatus Herbst, 1779 : 316; pl. 7, fig. 1. [Synonymized with atomarius Fabricius by
Panzer, 1801 : I.]

Elater carbonarius Schrank, 1781 : 184. [Synonymized with punctatus Herbst by Herbst,
1784 : I10.]

Elater pulverulentus Panzer, 1795 : 235. [Synonymized with atomarius Fabricius by Panzer,
1801 : I.]

Elater atomarius Fabricius, 1798 : 139.

Adelocera punctatus (Herbst) Seidlitz, 1888 : 168.

Lacon punctatus (Herbst) Fleutiaux, 1926 : 94.

Elater punctatus Herbst. Herbst appears to have based his description on material
in his own collection (1779 : 315) which is now preserved in the MNHN, Berlin (not
examined). Herbst does not record a locality but comments that punctatus is the
largest of the ‘einheimischen’ [native, presumably German] Elaterids.

The figure clearly depicts the distinctive species standing as punctatus Herbst in
all the collections known to me.

Elater carbonarius Schrank. The description is based on an unrecorded number of
specimens from Vienna. The location of the Schrank collection or of the specimens
on which he based his description is unknown. The description is sufficiently good
to allow Herbst’s synonymy to be accepted without reservation.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 77

Elater pulverulentus Panzer. Panzer does not make it clear whether his descrip-
tion of Elater pulverulentus is based on specimens in his own or some other collection
or on Herbst’s figure of Elater punctatus. Panzer is known to have possessed a
collection of insects (see obituary, 1829, Flora, Jena 12 : 400) which was probably
sold by auction after his death. Unfortunately the fate of the Coleoptera collection
is unknown. In the absence of any authentic material, Herbst’s figure of Elater
punctatus is here designated as the lectotype.

LECTOTYPE (present designation): Herbst, 1779: pl. 7, fig. 1 depicting Elater
punctatus Herbst, 1779 : 316.

Why Panzer did not use Herbst’s name is unknown. Arnett (1953 : 6) suggests
that the fact that Panzer refers only to Herbst’s figure indicates that he believed that
the figure and the description do not agree. Since Panzer does not refer to the
description elsewhere and also refers only (1779 : 200 and 227) to Herbst’s figures
in the case of Hispa atra and Buprestis berolinensis I do not believe this to be the
case. It seems more probable that Panzer merely wished to provide his readers
with a reference to a good figure illustrating his description, and considered Herbst’s
description to be unimportant. One possible reason why Panzer substituted the
name pulverulentus (powdery, dusty) for punctatus (spotted) is that he considered the
former to be a more accurate description of the species than the latter. Another
possibility is that he did so to avoid possible confusion.

Some time before the publication of the ‘Taschenbuch’ in 1795, Panzer
(1793 : 118) used the name Elater punctatus in his translation of Voet (1781 or
possibly earlier, see references p. 297). Panzer’s annotations indicate that he
considered Elater punctatus of Voet to be synonymous with Elater bipustulatus
Linnaeus but he may have believed that the name used in Voet’s attractive
illustrated work would have a wider circulation and be more readily accepted than
that published in Herbst’s paper, which appeared in the journal of a learned society.

Whatever the reason for his action, six years later (1801 : 1) he placed pulverulentus
Panzer, carbonarius Schrank and punctatus Herbst in synonymy with Elater
atomarius Fabricius. The available evidence suggests that this synonymy is fully
justified. Why Panzer gave preference to the Fabrician name is unknown. It
may have been because he wished to express his high esteem for Fabricius (Panzer,
1795, Vorbericht). Herbst (1806 : 14) accepted the use of atomarius Fabricius in
place of his own punctatus without comment.

Elater atomarius Fabricius. Fabricius credits this species to Linnaeus [1767]
Syst. Nat. 2. 655.28 and Fn. Sv. [1761 : 208] 738. However as Panzer (1801 : 1)
points out, Linnaeus did not describe a species with this name. Both references
refer to murinus Linnaeus. Fabricius lists E. pulverulentus Panzer as a synonym
and his comment ‘Habitat in Germania. Dom Panzer’ suggests that he may have
based his description of atomarius on Panzer’s pulverulentus specimens. Though
it can never be proved conclusively that pulverulentus Panzer and atomarius
Fabricius are objective synonyms, there seems no doubt that both authors were
referring to the same species and that Panzer’s synonymy can be accepted without
reservation.

78 Ci M. F. von HAYEK

Why Fabricius listed the references to murinus under atomarius remains a mystery.
He lists the same two references under murinus in the main body of the work (1792 :
221) though with a different, obviously erroneous, page number [22 instead of 28]
for the 1767 publication. Why he re-named pulverulentus Panzer is also unknown.
Arnett (1953 : 6) suggests that Fabricius did so because he was aware that pulveru-
lentus Panzer was a homonym of Elater pulverulentus Herbst, 1786 [= Chalco-
lepidius porcatus (Linnaeus)|. This explanation is unacceptable as Fabricius does
not refer to E. pulverulentus Herbst in any of his works and was probably unaware
of its existence. A more probable explanation is that he had already named a
specimen, and intended to publish the description of, Elater pulverulentus from
‘America meridionali’ in the Sehestedt collection. The description of this species,
which does not appear in the Schenkling catalogue, was published in 1801 (see
L. laticollis (Candéze) p. 68).

With the exception of a few workers who used the name carbonarius Schrank, the
name atomarius Fabricius appears to have been generally accepted for this distinctive
species until Seidlitz (1888 : 68) corrected the synonymy.

Lacon pyrsolepis (LeConte) comb. rev.

Adeloceva pyrsolepis LeConte, 1866 : 389.
Lacon pyrsolepis (LeConte) Fleutiaux, 1926 : 93.
Diphyaulon pyrsolepis (LeConte) Arnett, 1969 : II.

Holotype. U.S.A.: g, N.M.; Adelocera pyrsolepis Lec. [Ulke] (CM, Pittsburgh,
Ulke collection).

The published locality is New Mexico. The specimen agrees so well with the
description that despite the absence of LeConte’s determination label I have no
hesitation in accepting it as the holotype.

Lacon quadrinodatus Lewis

Lacon quadrinodatus Lewis, 1894 : 28.
Lacon (Kobulacon) quadrinodatus Lewis; Chaijo & Ohira, 1965 : 2.

Holotype. JAPAN: 9, Yuyama Chio 1883 [sic, on underside of card mount];
Japan, G. Lewis 1910.320; Yuyama [sic]; Lacon 4-nodatus Lewis, type [Lewis];
G. Lacon ex Adelocera [? Fleut.] (BMNH).

The published locality is Oyayama.

Lacon querceus (Herbst) comb. rev.

Elater quercea Herbst, 1784 : 113; pl. 27, fig. 11.

Elatey quercinus Gmelin, 1890 : 1912. [Unjustified emendation. ]

Elatey varius Olivier, 1790: no. 31, p. 32; no. 31, pl. 3, fig. 26, pl. 7, fig. 68.
Adeloceva quercus Seidlitz, 1872 : 110. [Unjustified emendation.]
Adelocera queyca Reitter, 1911 : 213. [Unjustified emendation.]

Lacon quercea (Herbst) Fleutiaux, 1926 : 94.

Zalepia quercea (Herbst) Arnett, 1959 : II.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 79

Elater quercea Herbst. The description is based on an unrecorded number of
specimens from Reppen [East Germany] and Hungary.

Type-material. Herbst’s material is preserved in the NMHU, Berlin (not
examined).

Examination of the material in the BMNH and the MNHN, Paris and other
collections has shown that there is no difference of opinion concerning the identity
of this species.

Elater varius Olivier. LECTOTYPE (present designation). FRANCE: sex
undetermined, Museum Paris, Bosc 1828; varia Fab.; type de Olivier [Fleut.];
varia Fab. France (MNHN, Paris). The abdomen is missing.

The published locality is ‘aux environs de Paris. Du Cabinet de M. Bosc.’

Olivier lists Elater querceus Fuesl. (=Herbst, 1784 in Fuesly] Archiv. ins. 5.
pag. 113, tab. 27, fig. 11 as a synonym of varius. Why Olivier considered it
necessary to rename Herbst’s species is unknown.

Many authors, including Herbst (1806: 36) himself and Candéze (1857 : 55),
used the name varius, crediting it to Fabricius (1792 : 222) with quercea Herbst
as a synonym. Fabricius (loc. cit.) credits the species to Olivier. Why.so many
nineteenth century workers did so remains a mystery. Perhaps it was to show
the great esteem in which they held Fabricius.

Lacon recticollis Fleutiaux
Lacon recticollis Fleutiaux, 1927 : 67.
LECTOTYPE (present designation). NortTH VIETNAM: 4, Hoa Binh, Lac Tho,
A. de Cooman; Lacon recticollis Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 2 9, 1 g, same locality labels as the lectotype but without
Fleutiaux’s determination labels (MNHN, Paris).

Lacon robustus (Fleutiaux)

Adelocera robusta Fleutiaux, 1902b : 213.
Lacon robustus (Fleutiaux) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). Himaraya: 9, Himalaya; Adelocera
robusta Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 29, same locality labels as the lectotype but without Fleutiaux’s
determination labels (MNHN, Paris).

The published locality is ‘Himalaya’ without further details. As there are
no subsequent records of the species the country of origin remains unknown.

Lacon rorulenta (LeConte) comb. rev.

Adelocera vorulenta LeConte, 1859b : 283.
Lacon rorulenta (LeConte) Fleutiaux, 1926 : 93.
Diphyaulon rorulenta (LeConte) Arnett, 1969 : II.

80 C. M. F. von HAYEK

Holotype. U.S.A.: 2, blue paper disk [=Washington, see p. 277]; A. rorulenta
LeC., aurorata LeC. [LeC.]; Type 2375 (MCZ, Harvard).

The published locality is Steilacoom [Washington]. The inclusion of the name
aurorata LeC. on the determination label presumably refers to LeConte’s (1857 : 18)
misidentification of the species.

Lacon ruber (Perty)

Elater ruber Perty, 1830 : 20; pl. 5, fig. 1.
Lacon (Monocyrton) rubra [sic] (Perty); Golbach, 19696 : 157.

The description is based on an unrecorded number of specimens collected near
S. Paulo, BRAzIL by Spix and Martius.

Type-material: Spix and Martius material in ZSBS, Munich according to
Horn & Kahle (1936 : 208). Not confirmed.

The generic attribution is based on the following specimen: g, Fry. Rio Jan°;
Adelocera rubra Perty, brasiliensis Cast., Brasilia [Fry] (BMNH).

This is the species generally known as rubra. This erroneous spelling was first
used by Candéze (1857 : 67).

Germar (1840) placed Adelocera brasiliensis Castelnau (1836: 14), described
from an unrecorded number of specimens from Brazil, in synonymy with ruber.
As the type material (?7NMV, Melbourne or IRSNB, Brussels) has not been located
the synonymy has not been confirmed.

Lacon rubidus (Schwarz)

Adeloceva vubida Schwarz, 1902b : 195.
Lacon (Aulacon) rubida (Schwarz); Golbach, 1969 : 157.

The description is based on an unrecorded number of specimens from Insula
Mona [PUERTO Rico].

Type-material: >DEI, Eberswalde.
The generic attribution is based on the description.

Lacon rubripennis Fleutiaux
Lacon (Adelocera des Auteurs) rubvipennis Fleutiaux, 19404 : 89.

LECTOTYPE (present designation). CurNa: 9, Tali [sic], Haut Yunnan; Lacon
rubripennis Fleut., type (Adelocera) [Fleut] (MNHN, Paris).
The published locality is Toli.

Lacon sachalinensis (Miwa) comb. n.
Adeloceva sachalinensis Miwa, 1927 : 13; pl. 1, fig. 1.

Holotype. ?CurINnA: Saghalien, collected by S. Issiki at Nairo, Aug. 10, 1914.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 81

The location of type not recorded. ?Taiwan Agricultural Research Institute,
Taipei.
The generic attribution is based on the description and figure.

Lacon salvazai (Fleutiaux)

Adeloceva salvazai Fleutiaux, 1918d : 184.
Lacon salvazai (Fleutiaux) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). NortH VIETNAM: 9, Laos, Xieng Khouang,
20.[?]13.15, V. de Salvaza; Adelocera salvazai Fleutiaux, Type [Fleut.] (MNHN,
Paris).

Lacon sanguineus (Candéze) comb. n.
Scelisus sanguineus Candéze, 1863 : 327; pl. 5, fig. 1.

Holotype. 9, India or.; Janson coll. 1903.130; Scelisus sanguineus Cdz. type
[Cand.]; Scelisus sanguineus Cand. [Janson] (BMNH).

The published locality is ‘Hindoustan septentrional’. As the term ‘Hindoustan’
was used for both the whole of India and also for a more restricted region north
of the Vindhya mountains, the locality remains uncertain. There are no additional
records of this species. The discrepancy between the published locality and the
label probably arose when Janson relabelled the specimens.

The characteristics of this species, the type-species of Scelisws, are discussed

on p. 54.

Lacon setosus (Candéze)

Adelocera setosa Candéze, 1874 : 29.
Lacon (Aulacon) setosa (Candéze); Golbach, 19696 : 158.

LECTOTYPE (present designation). BraziL: g, Villanova, Bates coll.; Janson
coll. 1903 : 130; Adelocera setosa mihi [Janson, ‘mihi’ is crossed out] Candéze,
type [Gahan] (BMNH).

The absence of Candéze’s label is probably due to Janson (see p. 276). The
published locality is Villa Nova de Rainha.

Lacon sparsus (Candéze) comb. rev.

Adeloceva sparsa Candéze, 1865 : 6.
Lacon sparsus (Candéze) Fleutiaux, 1926 : 93.
Diphyaulon sparsus (Candéze) Arnett, 1969 : IT.

LECTOTYPE (present designation). U.S.A.: 3g, Adelocera sparsa n. sp. Cdz.
Californie, Sacramento, Schaum [Cand.]; Janson coll., ex Candéze; Lectotype
Adelocera sparsa Cand. [M. C. Lane, 1964] (BMNH).

The specimen also bears M. C. Lane’s label; Lectotype, Adelocera sparsa Cand.
This lectotype designation does not appear to have been published.

F

82 CoM. 1B. von EASE

The material from the Dohrn collection recorded by Candéze, if it survived the
war, should be in the IZPAN, Warsaw (see p. 274).

Lacon spurcus (Candéze) comb. rev.

Adelocera spurca Candéze, 1874 : 25.
Lacon spurcus (Candéze) Fleutiaux, 1926 : 93.
Sulcilacon spurcus (Candéze) Fleutiaux, 194Ic : 48.

LECTOTYPE (present designation). Laos: g, Laos; Mouhot, Janson coll.
1903 : 130; A. spurca Cdz. type [Cand.]; Adelocera spurca Cand. Laos (Mouhot)
[Gahan] (BMNH).

Paralectotype: 9, Laos, Mouhot; Janson coll. 1903 : 130; A. spurca Cdz. type
[Cand.] (BMNH).
The material in the Saunders collection recorded by Candéze has not been traced

(see p. 279).

Lacon strangulatus (Fleutiaux)

Adeloceva stvangulata Fleutiaux, 1906a : 199.
Lacon stvangulatas (Fleutiaux) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). MADAGASCAR: 9, Amber Geb.; strangulata
Fleut., type, Region Diego Suarez [Fleut.]; Bull. Soc. ent. Fr. 1906, p. 199 [Fleut].
(MNHN, Paris).

Lacon subcostatus (Candéze)

Adeloceva subcostata Candéze, 1857 : 69.

Adeloceva subcostata var. a Candéze, 1857 : 69.

Adeloceva subcostata vhr. a. albicans Chevrolat, 1867 : 596.

Lepidelatey misticius Mignot, 1969 [March]: 12. Syn.n.

Lacon (Aulacon) subcostata (Candéze); Golbach, 19696 [November] : 159.

Adelocera subcostata Candéze. LECTOTYPE (present designation). LESSER
ANTILLES, GUADELOUPE: 9, Collection Chevrolat; subcostata Cand. type [Fleut.];
A. subcostata Chevr. Ann. Soc. Ent. Fr. 1867, p. 596 [Fleut.]; Fleut., Ann. Fr. rgi1,
p- 246 [Fleut.]; Adelocera subcostata Cand. Mon. 1, 1857, p. 69, 24. Guadeloupe,
Pointe a Pitre. L. Leherminier [Chevrolat, green label. The words ‘Cuba P.
Poey’ also appear squeezed in among the other words. They may have been
added later] (MNHN, Paris).

Candéze states that he retained the name under which Chevrolat submitted
the specimens. Chevrolat (1867 : 596) records that he had two ‘types’. The
second specimen has not been located.

Adelocera subcosta var. a. Candéze. LECTOTYPE (present designation).
CuBA: 9, Cuba, Collection Chevrolat; subcosta v. a. (albicans) Chevr. type [Fleut.]
(MNHN, Paris).

Lepidelater misticuus Mignot. Holotype. VirGin IsLanps: 3g, St John,

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 83

November 15, 1966 (in insect trap). USDA. lot 66.2777 (USNM, Washington).
Not examined.

Paratypes examined: 1 g, Isla Verde, Puerto Rico, 9.20.1968; in light trap.
V. L. Blackburn 68.24700; Paratype Lepidelater misticius Mignot 1 g, St John,
Virgin Is; G. Tallia, 8.17.67, 67.22778; Paratype, Lepidelater misticius Mignot
[aedeagus missing] (USNM, Washington). The location of the remaining 3 ¢
and I paratypes are unknown to me.

Mignot states that there is no evident secondary sexual dimorphism. In the
BMNH series of specimens the antennae of the males exceed the length of the
antennal groove by two segments, while in the female the last segment of the
antenna barely attains the end of the groove.

Lacon subpectinatus (Schwarz) comb. rev.

Alaotypus subpectinatus Schwarz, 1902f : 308.

Adeloceva denticornis Fleutiaux, 1906) : 212. [Synonymized by Fleutiaux, 1918d : 186.]
Lacon subpectinatus (Schwarz) Fleutiaux, 1926 : 93.

Alaotypus subpectinatus (Schwarz); Fleutiaux, 1927 : 62.

Alaotypus subpectinatus Schwarz: the description is based on two specimens
from N. VieTNAmM: Tonkin, Montes Mauson (Fruhstorfer).

Syntypes: ? DEI, Eberwalde.

Adelocera denticorms Fleutiaux. LECTOTYPE (present designation). NorTH
VIETNAM: 4, Tonkin Montes Mauson, Avril—Mai, 2-300, H. Fruhstorfer; Adelocera
denticornis Fleut., type [Fleut.]; Bull. Soc. Ent. Fr. 1906, p. 212 [Fleut.] (MNHN,
Paris).

Paralectotypes: 1 9, 2 g, same locality labels as the lectotype, but without
Fleutiaux’s determination labels (MNHN, Paris).

As the descriptions agree and both species are based on material from the same
locality, it seems reasonable to accept Fleutiaux’s synonymy.

Lacon tenebrionoides (Candéze)

Adelocera tenebrionoides Candéze, 1857 : 61.
Adelocera tenebrioides Candéze, 1874 : 22. [Unjustified emendation.]

LECTOTYPE (present designation). Mapacascar: 9, Laf. Madagascar; Janson
coll. 1903: 130; Adelocera tenebrionoides Cdze, Cand., type (e coll. Laferte
[Janson]; n. sp. Madagascar (BMNH).

The material from the Deyrolle collection recorded by Candéze has not been
traced.

Lacon tonkinensis (Fleutiaux) comb. n.
Alaotypus tonkinenis Fleutiaux, 1927 : 62; pl. 2, fig. 39.
LECTOTYPE (present designation). NortH VIETNAM: 9, Tonkin Chapa.

84 Cy M: EF. von HAYEK

Mai 1916. Vitalis de Salvaza; Adelocera tonkinensis Fleut., type [Fleut.]; type
de la fig. (MNHN, Paris).

Paralectotype: I ex., Tonkin, Chapa. 4.v.1918. Jeanvoine, without Fleutiaux’s
determination label (MNHN, Paris).

Lacon tumulosus (Candéze)

Adeloceva tumulosa Candéze, 1888 : 668.
Lacon tumulosus (Candéze) Fleutiaux, 1926 : 93.

Holotype. Burma: Plapoo, Tenasserim, aprile 1887, legit L. Fea. (MCZN,
Genoa, according to Binaghi, 1941c : 120.) Not confirmed.

The generic attribution is based on the description.

Lacon unicolor (Candéze)

Adelocera unicoloy Candéze, 1874 : 21.
Lacon unicoloy (Candéze) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). IRAN: 9, Persia; unicolor Cdz. [Cand.];
Adelocera unicolor Cand., type [Gahan] (BMNH).

Lacon vicinus Candéze

Adelocera vicina Candéze, 1891Ic : 772.
Lacon vicinus (Candéze) Binaghi, 1941c : 119.

The description is based on two specimens from BurRMA: Palon (Pégou) collected
ix.1888 by Fea.

Type-material. Burma: 1 4, Palon (Pegu) agosto-settembre 1877 [sic], legit
L. Fea (MCSN, Genoa according to Binaghi, 1941c : 120). Binaghi refers to this
specimen as the holotype but as the description is based on two specimens it must
be regarded as a syntype until a lectotype has been designated. The second
specimen may also be in the MCSN, Genoa or in the IRSNB, Brussels.

Candéze remarks that this species is probably a variety or subspecies of A. modesta
[Boisduval]. Fleutiaux (1926: 93) lists it as a synonym of Lacon modestus
(Boisduval), but Binaghi considers that it should be treated as a valid species.

The generic attribution is based on Binaghi’s redescription of the syntype.

Lacon viettei Girard
Lacon vietter Girard, 1970 : 25.

Holotype. Mapacascar: g, masif du Tsaratanana (versant sud), Andohana-
nahila, 1,850 m, III. 1967 (P. Soga) (MNHN, Paris).

Paratype: same data as holotype (Girard collection, Laboratoire de Zoologie,
Ecole normale supérieure, Paris).

Both specimens bear Girard’s determination labels.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 85

Lacon viridis (Champion)

Adelocera viridis Champion, 1894 : 260; pl. 11, fig. 3.
Lacon (Cornilacon) viridis (Champion); Golbach, 1969) : 158.

Holotype. Mexico: 9, Omitterne, Guerrero, 8,oooft. Aug. H. H. Smith;
B.C.A. Col. III (1); Adelocera viridis Ch. [Champ.]; sp. figured (BMNH).

Lacon vitalisi (Fleutiaux)

Adelocera vitalisi Fleutiaux, 1918d : 184.
Lacon vitalisi (Fleutiaux) Fleutiaux, 1926 : 93.

LECTOTYPE (present designation). NorTH VIETNAM: 9, Laos, Xieng Khouang
22.3.1915. R. V. de Salvaza; Adelocera vitalisi Fleut., type [Fleut.] (MNHN,
Paris).

Paralectotype: 2, Xieng Khouang 23.12.15, Vitalis de Salvaza (MNHN, Paris).

Lacon wallacei (Candéze)

Adeloceva wallacei Candéze, 1874 : 26.
Lacon wallacei (Candéze) Fleutiaux, 1926 : 93.

Holotype. Borneo: 9, Sar. 1363; Janson coll. 1903.130; Adelocera wallacei
(Janson) type [Janson] ms., Candéze [Gahan] (BMNH).
The absence of Candéze’s determination label is probably due to Janson (see

p. 276).

Lacon yunnanus Fleutiaux

Lacon (Adelocera des auteurs) yunnanus Fleutiaux, 19404 : 90.

LECTOTYPE (present designation). CuHina: 9, Kouang Si Hien, alt. 2109 m.
Sud Est Yunnan; Lacon (Adelocera) yunnanus Fleut. [Fleut.] (MNHN, Paris).

CANDANIUS nom. n.
Anius Candéze, 1889 : 103. Type-species: Anius gracillimus Candéze, by monotypy.

Anis Candéze, 1889 is a junior homonym of Anius Pascoe, 1885 (Coleoptera,
Curculionidae). Candanius is here proposed as a replacement name for Anius
Candéze nec Pascoe.

GENERIC DIaGNosIs. As for Lacon. The difference between the two genera is that in
Candanius the posterior prosternal process is turned upwards immediately behind the anterior
coxae. This characteristic is not generally of any importance at the generic level within the
Agrypninae and further studies may show that there is no justification for the retention of
Candanius as a separate genus (see also below).

86 CoM. EF. von HAW EK

Candanius gracillimus (Candéze) comb. n.

Anitus gracillimus Candéze, 1889 : 103.

The description is based on an unrecorded number of specimens from Chile
submitted by Fairmaire. Fleutiaux (1907: 170) described the three specimens
in the Candéze collection in the IRSNB, Brussels in some detail. He tentatively,
but correctly, identified them as two males and one female but expressed some
doubt that they belonged to the same species. I believe that they may be
conspecific. As Fleutiaux remarks, it is impossible to tell which specimen formed
the basis of Candéze’s description. For the present I consider that all three
specimens should be treated as syntypes and they have been labelled to this effect.

Syntypes examined. CHILE: 9, Anius n.g. gracillimus Cand. Chile [?Fairm.];
ex coll. Fairmaire; Collection E. Candéze; Anius gracillimus Cand. 9? Fleutiaux
det. 1903 [Fleut.]; Anius gracillimus Cd. det. E. Candéze [IRSNB curatorial label].
I g, Quillota; Collection E. Candéze; G.n. Anius Cdz. [Cand., green border]; n. sp.
gracillimus Cdz., Chile, Frm. [Cand., green border]; Anius gracillimus Cd. det.
E. Candéze [IRSNB curatorial label]. 1 3, Quillota; Collection E. Candéze;
Anus gracillimus Cand. ?g, Fleutiaux det. 1903 [Fleut.]; Chile, ex coll. Fairmaire;
Anius gracillimus Cd. det. E. Candéze [IRSNB curatorial label] (IRSNB, Brussels).

Candéze placed Anius in the sub-tribe Pomachilites near Psilonicus. However
all three specimens differ from Psilonicus and its allies in that tibial spurs are
absent, the claws bear setae at the base and the prosternopleural suture is grooved
for a greater (9, groove occupies about half the length of the suture) or lesser
(g, groove occupies slightly less than one third of the length of the suture) part
of its length for the accommodation of the antennae. I believe that the presence
of these characters fully justifies the inclusion of gracillimus in the Agrypninae.
Fleutiaux (1907 : 170) was of the same opinion as he considered that the genus
should be placed near Adelocera [sensu auct. nec Latreille, =Lacon Castelnau
of the present work].

In general appearance and the possession of short antennal grooves C. gracillimus
bears a distinct resemblance to the chilean Dilobitarsus vitticollis (Fairmaire and
Germain) and D. crux (Philippi). It differs from these species in that there are
no lobes on the tarsi. It is of interest that C. gracillimus resembles D. vitticollis
and crux in that the prothorax of the female is much more strongly convex than
that of the male and I consider that this lends support to my belief that the
syntype specimens are conspecific. It must be admitted that in no other species
known to me does the length of the antennal groove of the female differ from that
of the male. Fleutiaux’s (1907: 167) comment that this difference occurs in
vitticollis is based on the erroneous synonomy of D. vitticollis and D. crux.

For the present the genus Candanius is retained as a separate entity. Study
of additional material, when it becomes available, may show that (as in the case
of Agrypnus) the structure of the tarsi and the length and depth of the groove
for the accommodation of the antennae are not characters of sufficient impor-
tance to justify the retention of Dilobitarsus, Lacon and Candanius as separate
genera.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 87

DANOSOMA Thomson gen. rev.

Danosoma Thomson, 1859 : 103. Type-species: Elater conspersus Gyllenhal, by monotypy.
Delox Quelle, 1932 : 208 [as a subgenus of Adelocerva sensu auct., nec Latreille]. Type-species:
Elater conspersa Gyllenhal, by subsequent designation (Arnett, 1955 : 607).

GENERIC DIAGNOSIS. Tarsal claws without a group of setae at the base (Text-fig. 12).
Tibial spurs absent (Text-fig. 10). Mesepisternum does not from part of the margin of the
mesocoxal cavity; mesepimeron forms part of margin of mesocoxal cavity (Text-fig. 3). Second
antennal segment moniliform, third weakly triangular, each almost equal in length to the fourth
and following segments. Antennal grooves long, extending almost the entire length of the
prosternopleural suture and deep enough to accommodate the antennae. Body clothed with
narrow scales. Prothorax simple, without constriction behind the anterior angles; lateral
carina present. Propleurae and metasternum without well-defined grooves for the accommoda-
tion of the tarsi of the anterior and middle legs. Scutellum simple, without carina. Tarsi
simple, without ventral lobes (Text-fig. 15).

RANGE OF VARIATION FOUND WITHIN THE GENUS. The four species included
in the genus bear a close resemblance to one another and also to certain Lacon
species from which they are easily distinguished by the absence of setae at the base
of the claws. The curious similarities between certain palaearctic and holarctic
species are discussed in the section on the distribution of the species.

HisTORY OF THE GENUS. Thomson established Danosoma for a single species,
conspersa Gyllenhal, in which the antennal grooves are long enough to accommodate
the antennae and the base of the prothorax is narrower than the base of the elytra.
In addition the posterior angles of the prothorax are bicarinate and not turned
outward at the apex. Seidlitz (1888 : 169) reduced Danosoma to a subgenus of
Adelocera (sensu auct. nec Latreille = Lacon Castelnau of the present work).
The subgenus was not generally accepted and Arnett (1953 : 7) treated Danosoma
as a subjective synonym of Lacon Castelnau.

Quelle was aware of the existence of Danosoma but nevertheless established
Delox as a subgenus of Adelocera (sensu auctt., nec Latreille) for four species,
including conspersa Gyllenhal, which lacked setae at the base of the claws.
Arnett (1955 : 607) designated conspersa as the type-species of Delox, thereby
making Danosoma and Delox isogenotypic.

I consider the absence of the setae at the base of the claws to be of sufficient
importance to warrant the establishment of a genus. The genus Danosoma is
here redefined on this basis.

SEXUAL DIMORPHISM. The species included in this genus do not appear to
display any marked sexual dimorphism.

DistrisuTion. Northern Palaearctic and northern Holarctic regions. Two
species occur in each region. There is a close resemblance, both in general
appearance and in the shape of the aedeagus, between the palaearctic conspersa
and the holarctic brevicornis and the palaearctic fasciata and the holarctic obtectus.

Quelle (1932: 209) drew attention to the similarities between conspersa and
brevicornis and fasciata and his interpretation of profusa. Quelle misinterpreted
profusa, believing that the posterior angles of the prothorax were simple, whereas

88 Ce. M. Fs yon HAW EK

those of the type are laterally compressed. Quelle’s profwsa [not examined] is
probably obtectus Say.

BIOLOGY AND HABITS. The larvae of D. fasciata (Linnaeus) has been described
by Perris (1876: 169). The larvae are predaceous and live in wood, probably
in the galleries of wood-boring insects (Kirk, 1933).

SPECIES INCLUDED IN THE GENUS

Four species are at present assigned to the genus. Three of these were included
in Delox by Quelle. The fourth, obtectus Say, is probably the species which was
known to Quelle as profusa.

Danosoma brevicornis (LeConte) comb. n.

Adeloceva brevicornis LeConte, 1853 : 491.

Adelocera profusa Candeze, 1857 : 54. [Synonymized by Arnett, 1952 : 115.]

Adelocera vetusta Walker, 1866 : 324. [Synonymized with profusa by Candeze, 1891Ic : 13.]
Adelocera (Delox) brevicornis (LeConte); Quelle, 1932 : 208.

Lepidotus (Danosoma) brevicorms (LeConte) Arnett, 1952 : I15.

Lacon (Lacon) brevicornis (LeConte); Arnett, 1953 : 7.

Adelocera brevicornis LeConte. LECTOTYPE (present designation). U.S.A.:
9, light blue-grey paper disk [=Lake Superior, see p. 277]; Type 2377 [MCZ
curatorial label]; Ad. brevicornis LeC. Lac. Sup. [LeConte] (MCZ, Harvard).

Paratypes: I g, 2 9, all with light blue-grey paper disks and labelled “brevicornis
4, 3 and 2 respectively (MCZ, Harvard).

Adelocera profusa Candéze. Holotype. U.S.A.: g, Laf. Oregon; Janson coll.
1903.130; Oregon. Coll. Reiche; A. profusus Cdz., type [Cand.]; Holotype
Adelocera profusa Cand. M. C. Lane 1964 [M. C. Lane]; Adelocera profusa Cdze,
Cand. type (ex coll. de Laferté) [Janson] (BMNH).

Adelocera vetusta Walker. LECTOTYPE (present designation). CANADA: dg,
Brit. Colum./64.18 [BMNH registration number = presented by J. K. Lord];
vetusta [Walker]; Adelocera vetusta Walker, Brit. Columbia (J. K. Lord) [Gahan];
holotype Adelocera vetusta Walk., M. C. Lane 1964 [Lane] (BMNH).

Candéze (1891c : 13) lists Adelocera cavicollis LeConte, 1859 : 86 as a synonym.
The description is based on a single specimen found on the shores of Tlamath Lake
[? Upper Kalmath Lake, Oregon, Arnett, 1952: 115] by Dr Newberry. Arnett
(1952 : 115) states that this specimen is not in the LeConte collection. I have
been unable to trace Dr Newberry’s collection.

Arnett (1952: 115) believes that operculatus Say, 1836, nomen nudum, may
belong to this species.

Danosoma conspersa (Gyllenhal) comb. rev.

Elater conspersa Gyllenhal, 1808 : 377.
Danosoma conspervsa (Gyllenhal) Thomson, 1859 : 103.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 89

Adelocera (Delox) conspersa (Gyllenhal); Quelle, 1932 : 208.
Lepidotus (Danosoma) conspersa (Gyllenhal) Arnett, 1952 : 115.
Lacon (Lacon) conspersa (Gyllenhal); Arnett, 1953 : 7.

The description is based on an unrecorded number of specimens from SWEDEN.
Type-material: ? ZUM, Uppsala.

The generic attribution is based on the examination of specimens identified as
conspersa in the BMNH and MNHN, Paris.

Tscherepanov (1957: 190) includes conspersa in Adelocera. This is probably
due to the fact that he was unaware of Fleutiaux’s (1926) correction to the
Schenkling Catalogue in which the species is transferred to Lacon.

Danosoma fasciata (Linnaeus) comb. n.

Elater fasciatus Linnaeus, 1758 : 406.

Adeloceva angustata Sahlberg, 1903 : 25. [Junior primary homonym of Adelocera angustata
Philippi, 1861. See p. 100.}

Adelocera sahlbergi Schwarz, 1907 : 315. [Replacement name for Adelocera angustata Sahlberg.
Synonymized by Tscherepanovy, 1957 : 187.]

Adeloceva (Delox) fasciata (Linnaeus); Quelle, 1932 : 208.

Elater fasciatus Linnaeus. LECTOTYPE (present designation): 18, fasciatus
[Linnaeus] (BMNH, Linnaean coll.).

Paralectotypes: I ex., 27 [printed]. 1 ex., 95 [MSS.] (BMNH, Linnaean coll.).

Adelocera angustata Sahlberg. LECTOTYPE (present designation). U.S.S.R.:
g, Jeniseisk [sic]; J. Sahlb., spec. type; Adelocera angustata J. Sahlb. [Sahlb.]
(TU, Finland).

Paralectotypes: 1 g, Jeniseisk, Kitmanoff [sic] (TU, Finland). 1 3, 392 [museum
catalogue number = River Ob, between Narym and the mouth of the R. Tschulin,
I-2 June]; Mus. Zool. H.fors. spec. type No. 1222, Adelocera angustata J. Sahlberg;
A. angustata [?Sahlb]. The specimen also bears two labels, ‘Krasnojask’ and
‘Stalloff? which are believed to have been affixed in error (ZMU, Helsinki).

The published localities of angustata are: Habitat in Siberia. Prope oppidum
Timsk ad flumen Obi d. 1 Juni a Dom Trybom detecta; deinde prope oppidum
Jenisseisk d. 14 juni nonnulla specimina ipse inveni; postea etiam a Dom. Kipmanov
e vicinitate oppidi Jenisseisk missa.

Timsk is probably Tymsk or Ust’Tym (59°29’N., 80°03’E.) on the River Ob
about 300 km N.W. of the confluence of the Ob and Tschulin [Chulym] rivers.
Narym (59°00’N., 81°30’E.) is also on the River Ob about 200km N.W. of the
confluence of the Ob and Tschulin. Despite the discrepancy between the published

locality and that on the specimen I believe that it formed part of Sahlberg’s original
series.

Danosoma obtecta (Say) comb. n.

Elatey obtectus Say, 1839 : 181.
Lepidotus (Danosoma) obtectus (Say) Arnett, 1953 : 7.
Lacon (Lacon) obtectus (Say); Arnett, 1969 : 11.

go CM. EFS von EOAY EK

The description is based on an unrecorded number of specimens from
MASSACHUSETTS submitted by Dr Harris.

Type-material: not found in the ANS, Philadelphia, see note on the Say
collection, p. 279. There are no specimens of this species in the Harris collection
in the MCZ, Harvard. The type-material must therefore be regarded as lost.

The generic attribution is based on the following specimen standing as obtecta
Say in the BMNH: 3, Buffalo, N.Y.; Andreas Bolter Collection.

The description of this species may have appeared before 1839, see note in the
bibliography, p. 295.

EIDOLUS Candéze
Exidolus Candéze, 1857 : 178. Type-species: Eidolus linearis Candéze, by monotypy.

GENERIC DIAGNOSIS. Each claw with a group of setae near the base (Text-fig. 11). Tibial
spurs absent (Text-fig. 10). Mesepisternum does not form part of margin of the mesocoxal
cavity; mesepimeron forms part of margin of the mesocoxal cavity (Text-fig. 3). Third antennal
segment larger than the second, similar in shape to the fourth. Segments three to eleven
pectinate. Antennal grooves shallow, occupying the anterior three-quarters of the
prosternopleural suture. Body clothed with narrow scales. Prothorax without constriction
behind the anterior angles; lateral carina absent. Propleurae and metasternum without
grooves or depressions for the accommodation of the anterior and middle tarsi. Scutellum
simple, without carina. Tarsi simple, without ventral lobes.

HIsTORY OF THE GENUS. The genus was established for a single species with
pectinate antennae in which the lateral margin of the prothorax is not carinate.

SEXUAL DIMORPHISM. Unknown. The two lectotypes are the only specimens
known to me.

DISTRIBUTION. Brazil.

LIFE HISTORY AND HABITS. Nothing is known of the life history and habits
of the two species included in the genus.

SPECIES INCLUDED IN THE GENUS

Only two species are attributed to the genus.

Eidolus gratiosa (Fleutiaux) comb. n.

Adelocera grvatiosa Fleutiaux, 19026 : 213.
Lacon (Cornilacon) gratiosa (Fleutiaux); Golbach, 19690 : 188.

LECTOTYPE (present designation). Brazit: 9, Jatahay, Goyas, Brazil;
Adelocera gratiosa Fleut. type Bull. Soc. Ent. Fr. 1902, p. 213 [Fleut.] (MNHN,
Paris).

Eidolus linearis Candéze
Eidolus linearis Candéze, 1857 : 179; pl. 3, fig. I.

LECTOTYPE (present designation). Brazit: g, Braz. mer.; Janson coll.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE gI

1903.130; Eidolus linearis Cdze, type. [Janson] ex coll. Laferte [Gahan]; Hem.
agrestis Buq. in mus. e coll. de Laferte [Janson] (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see
p. 276). The second syntype, from the Deyrolle collection, has not been located.

OPATELUS Candéze

Opatelus Candéze, 1857: 86. Type-species: Opatelus limbatus Candeze, by subsequent
designation (Hyslop, 1921 : 660).

GENERIC DIAGNOsIS. Each claw bearing a group of setae near the base (Text-fig. 11)
Tibial spurs absent. Mesepisternum, does not form part of the margin of the mesocoxal cavity;
mesepimeron forms part of the margin of the mesocoxal cavity (Text-fig. 3). Second and third
antennal segments subequal, each smaller than the fourth and following segments. Antennal
groove long, extending almost the entire length of the prosternopleural suture. The distance
separating the posterior end of the groove from the anterior coxae is equal to the width of the
groove. The groove is deep enough to accommodate the rolled antennae. Body clothed with
narrow scales. Prothorax simple, without constriction behind anterior angles; lateral carinae
present. Propleurae and metasternum with well defined grooves for the reception of the tarsi
of the anterior and middle legs. Propleural grooves parallel to antennal grooves; metasternal
grooves curved, running diagonally across the sternite, the posterior ends, if produced, would
cut the posterior margin of the metasternum near the posterior lateral angles (Text-fig. 4).
Scutellum simple, without carina. Fourth tarsal segment with a ventral lobe (Text-fig. 16).

HIsTORY OF THE GENUS. This distinctive and easily recognized genus was
established for the four species which are still assigned to it.

Candéze’s generic diagnosis, which except for the first three characters listed,
agrees remarkably well with that given above, defines the genus so accurately that
it is very surprising that he subsequently assigned argentatus (see Lacon argentatus
(Candéze), p. 56) to the genus.

SEXUAL DIMORPHISM. The sexes are not distinguishable on external
characteristics.

DISTRIBUTION. Brazil, Bolivia.

BIoLoGy AND HABITS. Nothing is known of the habits or life-history of the
four species included in the genus.

SPECIES INCLUDED IN THE GENUS

Schenkling (1925 : 20) records five species in this genus. The generic attribution
of three species is confirmed by examination of the type-material and a fourth
provisionally placed on information available from the description. The fifth,
argentatus Candéze is transferred to Lacon (see p. 56).

Opatelus cribratus (Blanchard)

Lacon cribvatus Blanchard, 1845 : 145.
Opatelus cribratus (Blanchard) Candéze, 1857 : 88.

LECTOTYPE (present designation). Bortvia: 9, Museum Paris, Bolivie

92 CAM. Bw VON TEVA ETS

(Chiquitos) D’Orbigny, 1834; L. cribratus Blanch., Chiquitos, Mr. d’Orbigny; Lacon
cribratus E. Blanch., type Fleutiaux det. [Fleut.] (MNHN, Paris).

The date of publication of this species, which is generally recorded as 1837-43,
was obtained from Sherborn and Griffin (1934).

Opatelus fasciatus Candéze
Opatelus fasciatus Candéze, 1857 : 89.

The description is based on a unique specimen from Brazil submitted by
Boheman.

Holotype: ? NR, Stockholm.
The generic attribution is based on the description.

Opatelus limbatus Candéze
Opatelus limbatus Candéze, 1857 : 87; pl. 3, fig. 2.

LECTOTYPE (present designation). Brazit: 9, Dej. Brazil; Janson coll.
1903.130; Opatelus limbatus Cdze, Cand. [Janson]; Agrypnus punctipennis Dej.
Cat. e coll. Dejean [Janson] (BMNH).

Paralectotypes. 2 3, 1 9, with the same labels as the lectotype (BMNH).

The absence of Candéze’s and Dejean’s labels is probably due to Janson (see

p. 276).

Opatelus paleolatus Candéze
Opatelus paleolatus Candéze, 1857 : 88.

Holotype. 3, Brazit, Deyr.; Janson coll. 1903.130; O. paleolatus [Cand.];
Opatelus paleolatus Cdze, Cand., type (e coll. Deyrolle) [Janson] (BMNH).

ACROCRYPTUS Candéze

Cryptotarsus R. A. Philippi, 1873 : 308. Type-species: Cryptotarvsus ater R. A. Philippi, by
monotypy. [Homonym of Cryptotarsus Kirsch, 1865 : 88 (Malachiidae). |

Acrocryptus Candéze, 1874 : 39. [Replacement name for Cryptotavsus RK. A. Philippi not
Kirsch. ]

Hexaulacus Candéze, 1784: 40. Type-species: Hexaulacus veedi Candéze by monotypy.
[Synonymized by Golbach, 1970b : 307.]

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae near the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepisternum does not form part of the margin of the
mesocoxal cavity; mesepimeron forms part of the margin of the mesocoxal cavity (Text-fig. 3).
Third antennal segment resembles the fourth; both are larger and of a different shape than the
second segment. Antennal grooves deep, almost attaining the anterior coxae; the distance
between the end of the groove and the coxal cavity equal to the width of the groove. Body
clothed with scales; scales on the underside of the body narrower than those of the upper surface.
Prothorax not constricted behind the anterior angles. Lateral carina of the prothorax attains

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 93

the anterior margin. Grooves for the reception of the anterior, middle and hind tarsae present
on the propleurae, metasternum and first visible abdominal sternite. Scutellum simple,
without carina. Second, third and fourth tarsal segments with small ventral lobes.

HISTORY OF THE GENUS. Cryptotarsus was established for a single species from
Chile with grooves on the propleurae for the reception of the anterior tarsi.
Hexaulacus was also established for a single Chilean species with grooves for the
reception of the anterior, middle and hind tarsi on the propleurae, metasternum
and first visible abdominal segment. Calvert (1889: 18) suggested that C. ater
Philippi and H. rveedi Candéze were conspecific but Fleutiaux was of the opinion
that this could not be the case as Philippi mentions only one pair of tarsal grooves.
Golbach (1970) : 306) examined the type of C. ater and found that six tarsal grooves
are present as in veedi. As Golbach suggests, the position of the middle and hind
legs probably prevented Philippi from seeing the tarsal grooves on the metasternum
and abdomen.

SPECIES INCLUDED IN THE GENUS

Acrocryptus ater (Philippi)

Cryptotarsus atey R. A. Philippi, 1873 : 308.
Acrocryptus ater (Philippi) Candéze, 1874 : 39.
Hexaulacus veedi Candéze, 1874 : 40. [Synonymized by Golbach, 1970b : 307.]

Acrocryptus ater Philippi. Holotype. CHILE: near Rancagua (MN, Santiago
according to Golbach, 1970) : 306). Not examined.

Hexaulacus reedi Candéze. Holotype. CHILE: 9, 53884 [Fry manuscript
catalogue number recording locality and determination]; Reed; Chile, Valda.;
Fry coll. 1905.100; Hexaulacus reedi Cdz. n. gen. n. sp. non Cryptotarsus ater
[Cand.]; Cryptotarsus ater Ch. MS. [illegible] Valdivia; Hexaulacus reedi Candéze;
Chile, type [Fry] (BMNH). Not lost as stated by Golbach (19700 : 307).

Rancagua is about four hundred miles north of Valdivia.
Nothing is known of the life-history or habits of this species.

DILOBITARSUS Latreille

Dilobitarsus Latreille, 1834 : 142. Type-species: Dilobitarsus tuberculatus Latreille [= Dilobi-
tarsus bidens (Fabricius)], by monotypy.

Anacantha Solier, 1851: 18. Type-species: Anacantha sulcicollis Solier, by monotypy.
Syn. n.

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae at the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepisternum does not form part of the margin of the
mesocoxal cavity; mesepimeron forms part of the margin of the mesocoxal cavity (Text-fig. 3).
Third antennal segment larger than the second, similar to the fourth. Antennal groove of
variable depth, always extending beyond the anterior half of the prosternopleural suture.
Body clothed with narrow scales. Prothorax not constricted behind the anterior angles; lateral
carina present. Pronotum with or without tubercles. Propleurae and metasternum without

94 CoeM. “By Vion EAE

well defined grooves for the reception of the anterior and middle tarsae. Scutellum simple,
without carina. Third and fourth tarsal segments with ventral lobes.

NOTES ON THE GENERIC SYNONYMY. Almost a hundred years ago Candéze
(1874 : 39) drew attention to the fact that Dzlobitarsus and Anacantha were not
readily separable. Examination of the available material has shown that the
three main characteristics used to separate the genera, the appearance of the
prothorax, the length of the antennae and the antennal grooves vary considerably
from one species to another. In my opinion the retention of the two genera
cannot be justified.

Fleutiaux (1907) : 168) retained the genus Anacantha using the appearance of
the pronotum as the diagnostic characteristic of the genus. He restricted the
genus to the type-species and transferred the remaining species at that time
assigned to the genus (vitticollis (Fairmaire & Germain), marmorata Candéze and —
fairmaiver Candéze to Adelocera (sensu auctt., nec Latreille = Lacon Castelnau
of the present work). In view of the known variability of the appearance of the
prothorax in the Elateridae and the fact that at the present time the known species
can be assigned to Dilobitarsus or Lacon on the basis of the structure of the tarsi,
I do not consider Fleutiaux’s action with regard to vitticollis and marmorata was
justified. A. fairmairvei is unknown to me. However it must be admitted that
certain species at present assigned to Dzulobitarsus or Lacon on the basis of the
structure of the tarsi bear a strong resemblance to one another, e.g. L. laticollis
(Candéze) and D. abbreviatus Candéze. See also the note on the structure of the
fourth tarsal segment of L. dubius Candéze (p. 63). It seems possible that, as
more material for study becomes available, the structure of the tarsi will be found
to vary from one species to another in a manner which precludes this characteristic
from being used to separate two genera.

HISTORY OF THE GENUS. The genus was established for a single Central American
species. In the years which followed a number of species from both Central and
South America and Africa were assigned to the genus. Schwarz (1902)) established
the genus Elasmosomus to accommodate the African species. With the exception
of Fleutiaux, workers have accepted Duzlobitarsus and Elasmosomus as separate
genera. Fleutiaux (191g : 11) continued to assign African species to Dulobitarsus
and may have advised Paulian (1947) to do the same when he described fleutiauxa
(p. 104). For Fleutiaux’s treatment of Dilobitarsus and Elasmosomus in 1941 see
Dp: 10m

SEXUAL DIMORPHISM. In swicicollis (Solier) and crux (Philippi) the males differ
from the females in that the body is more slender, the lateral margins of the
prothorax straight instead of arcuate and the antennae more strongly serrate and
longer, attaining the posterior angles of the prothorax. In the remaining species
known to me these differences are less apparent.

DISTRIBUTION. Central and South America. A single species, pendleburys
Fleutiaux from Borneo is also assigned to the genus.

BIOLOGY AND HABITS. Nothing is known of the life history and habits of
Dilobitarsus species.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 95

SPECIES INCLUDED IN THE GENUS

The generic attributions of nineteen of the twenty-three species listed as Dzlo-
bitarsus in the Schenkling catalogue (1925) or described subsequently have been
confirmed. D. cuneatus Candéze is transferred to Lacon (p. 62), D. minutus
Candéze to Adelocera (p. 39) and D. fleutiauxt Paulian (1947) to Elasmosomus
(p. 104). D. impressicollis Schwarz (1902d: 129) is unknown to me. Two of the
species removed from Anacantha by Fleutiaux (1907) : 168) (marmorata and
vitticollis) are assigned to Dilobitarsus. The third (fairmairei Candéze) is unknown
to me. Finally, crux Philippi is extracted from synonymy with vitticollis and
included in Dilobitarsus as a valid species.

Dilobitarsus abbreviatus Candéze
Dilobitarsus abbreviatus Candéze, 1857 : 78.
The description is based on a single unidentified Brazilian specimen from the
Germar collection submitted by Schaum.

Holotype. The holotype appears to have been lost. It cannot be found (see
notes on the Schaum and Candéze collections, p. 280 and p. 271) in the BMNH or
IRSNB, Brussels.

The generic attribution is based on the following specimen: 3, Fry, Rio Jano;
abbreviatus Cdz., Br. [Cand., green border] (IRSNB, Brussels).

Dilobitarsus bicornis Candéze
Dilobitarsus bicornis Candéze, 1857 : 82.
The description is based on an unrecorded number of specimens from BRAZIL
submitted by Riehl under the name D. tuberculatus Latreille.

Type-material: There are no specimens determined by Candéze in the BMNH.
?IRSNB, Brussels. The Riehl collection passed to the University Zoological
Museum, Marburg (Horn & Kahle, 1936 : 225). Not examined.

The generic attribution is based on the following specimen: 3, Rio de Janeiro.
| Dec. 1856. J. Gray; Dilobitarsus bicornis Cand. [Janson] (BMNH).

Dilobitarsus bidens (Fabricius)

Elater bidens Fabricius, 1801 : 227.
Dilobitarsus tuberculata Latreille, 1834 : 143 [Synonymized by Candéze, 1857 : 83.]
Dilobitarsus bidens (Fabricius) Candéze, 1857 : 83.

Elater bidens Fabricius. The description is based on an unrecorded number
of specimens from ‘America meridionali D. Smidt Mus. D. Lund’.

Type-material: 1 ex., ZMU, Copenhagen according to Zimsen (1964: 158).
Not examined.

96 CC. MS ES ‘vio EVAGY Bi

The interpretation of the species is based on specimens in the BMNH determined
by Candéze and other workers.

Dilobitarsus tuberculata Latreille. The description is based on an unrecorded
number of specimens approximately 6 lines [=13-5 mm] long from Brazit. This
material should be in the BMNH (see p. 277) but the most careful searches have
failed to discover any specimens with Latreille’s determination label, or indeed
any specimens bearing the name tuberculata. The specimen recorded below agrees
so well with the description that I believe it may be part of Latreille’s original
material.

The interpretation of the species is based on the following specimen: BRAZIL: 9,
Dej. Brazil; Brazilia Latreille [probably Latreille]; Janson coll. 1903.130; Agrypnus
macrothorax Dej. Cat., e coll. Dejean [Janson]; Dilobitarsus bidens F., Cdze.,
Cand. [Janson] (BMNH). Length: 15-8 mm.

The species is distinctive and there seems to be no good reason why Candéze’s
synonymy should not be accepted.

Dilobitarsus cariosus Candéze

Dilobitarsus cariosus Candéze, 1889 : 71 (5).

Holotype. COLOMBIA: 9, n.sp. cariosus Cdz. Col. Ptand [Cand., green border,
last word illegible]; type [Cand.]; Collection E. Candéze; Dilobitarsus cariosus
Cand. det E. Candéze [IRSNB curatorial label] (IRSNB, Brussels).

The published locality is ‘Nouvelle Grenade’, the name by which Colombia
was known between 1831 and 1861.

Dilobitarsus colombianus Candéze

Dilobitarsus colombianus Candéze, 1857 : 78.

LECTOTYPE (present designation). CoLompia: 9, Laf. Columbia [sic]; Janson
coll. 1903 : 130; Agrypnus colombianus Buq. Colombie, D. Jurgens [?Buquet];
Dilobitarsus columbianus [sic] Cdze, Cand., type e coll. de Laferté [ Janson)
(BMNH).

The published locality is ‘Nouvelle Grenade’ the name by which Colombia was
known between 1831 and 1861. The absence of Candéze’s determination is
probably due to Janson (see p. 276).

Dilobitarsus corrosus Candéze
Dilobitarsus covrosus Candéze, 1897 : 7.

LECTOTYPE (present designation). Botivia: 3, 1894, corrosus Cand. La Paz,
Bolivie St. [Cand., green border]; type [Cand.]; Collection E. Candéze; Dilobitarsus
corrosus Cand. det. E. Candéze [IRSNB curatorial label] (IRSNB, Brussels).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 97

Dilobitarsus crux (Philippi) sp. rev., comb. n.

Adelocera crux F. A. E. Philippi, 1860 : 247.

The description is based on an unrecorded number of specimens from ‘terra
Arancanorum’ [S. CHILE, between 37°29’S. and 40°18’S. and bounded by the
Pacific to the west and the Andes to the east].

Type-material: ?>MN, Santiago (Horn & Kahle, 1936 : 208).

The present interpretation of cyux is based on a number of specimens in the
BMNH collection from Chile (Reed) determined by Candéze. They agree well
with the description and differ from vitticollis (Fairmaire and Germain) (p. 100)
with which Fleutiaux (1907) : 166) synonymized the species, in the following
manner: The yellow scales on the pronotum are limited to the anterior and
posterior angles, so that the dark scales clothing the rest of the surface form a
well defined cross, the third antennal segment is distinctly shorter than the fourth
and the propleural antennal groove is shallow posteriorly. There is also a slight
but distinct difference in the shape of the tips of the lateral lobes of the aedeagus.

Dilobitarsus deyrollei Candéze
Dilobitarsus deyrollei Candéze, 1857 : 79.

Syntype examined: CoLtomsia: 3g, Laf. Columbia [sic]; Janson coll. 1903 : 130;
Agrypnus bituberculatus Buq. Colombie [?Buquet]; Dilobitarsus deyrollei Cdze,
type e coll. de Laferté [Janson] (BMNH).

The published locality is ‘Nouvelle Grenade’ [S. Colombia, see colombianus above].
The absence of Candéze’s determination label is probably due to Janson (see p. 276).
Since the second syntype was in the collection of Deyrolle (see p. 274), in whose
honour the species is named, the designation of the lectotype is postponed until
this specimen has been located.

Dilobitarsus eloini Candéze
Dilobitarsus eloini Candéze, 1874 : 35.
The description is based on an unrecorded number of specimens from CENTRAL
AMERICA.
Type-material: ?IRSNB, Brussels.

The generic attribution based on the following specimen, which may be a syntype:
2, Dilobitarsus eloini Cdz., Mexique [Cand.]; Dilobitarsus eloini Cand. Mexico,
type [Gahan]; coll. Janson, ex Candéze (BMNH).

Dilobitarsus gracilis Candéze
Dilobitarsus gracilis Candéze, 1874 : 35.
LECTOTYPE (present designation). BraziL: 9, Bates coll. Ega; Janson coll.
1903 : 130; Dilobitarsus gracilis mihi n.sp. [Janson, ‘mihi. n.sp.’ is crossed out

and replaced by ‘Candéze’ by Gahan] (BMNH).
G

98 Cc. M. F. von HAYEK

Paralectotype. BraziL: 9, Bates coll. Japajos [sic]; Janson coll. 1903 : 130;
Dilobitarsus gracilis mihi var. [ Janson, ‘mihi’ is crossed out and replaced by ‘Cand.’
by Gahan see p. 276] (BMNH). The published localities are R. Tapajos, Ega
and Para. The material from Para has not been traced.

Dilobitarsus inopinus Candéze
Dilobitarsus inopinus Candéze, 1874 : 35.

LECTOTYPE (present designation). NicaRAGua: 9, Nicaragua/EMJ; Janson
coll. 1903 : 130: Dilobitarsus inopinus mihi, type [Janson, ‘mihi’ is crossed out
and replaced with ‘Candéze’ by Gahan] (BMNH).

Dilobitarsus irroratus Candéze

Dilobitarsus ivvroratus Candéze, 1857 : 80.

LECTOTYPE (present designation). CoLompia: 9, Laf. N. Grenada; Nov.
Grenada, Goudot; Janson coll. 1903.130; Dilobitarsus irroratus Cdze., type e coll. de
Laferté [Janson] (BMNH).

Paralectotype. 9, Laf. N. Grenada; Janson coll. 1903.130; Dilobitarsus irroratus
Cdze, type e coll. de Laferté [ Janson] (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see
p- 276). N. Grenada = Colombia, see D. colombianus, p. 96.

Dilobitarsus laconoides (Fleutiaux) comb. n.

Anacantha marmorata Candéze, 1874 : 38.

Adelocera marmovata (Candéze) Fleutiaux, 1907) : 163. [Junior secondary homonym of
Adelocera marmorata (Fabricius, 1801).]

Adelocera laconoides Fleutiaux, 1907): 163. [Replacement name for Adelocera marmorata
(Candéze).]

Lacon (Cornilacon) laconoides (Fleutiaux) ; Golbach, 1969 : 158.

LECTOTYPE (present designation). CHILE: g, Chili, Reed; 53883 [Fry coll.
catalogue number = type from Reed coll.]; Dilobitarsus marmorata Cdz. s.n.
[Cand.] (BMNH).

Dilobitarsus lignarius Candéze
Dilobitarsus lignarius Candéze, 1857 : 80.

LECTOTYPE (present designation). BraziL: 9, Dej. B. Ayres. Braz.; Dilo-
bitarsus lignarius (Dej.) Cand. type, Cdze [Janson]; Agrypnus lignarius Dej. Cat.
e coll. Dejean [Janson] (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see

p: 276):

| aml

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 99

Dilobitarsus nebulosus Candéze
Dilobitarsus nebulosus Candéze, 1874 : 32.
Holotype. FRENCH GUIANA: 9, Cayenne; Janson coll. 1903 : 130; coll. Janson

ex. Candéze; ex. coll. Castelnau [Cand.]; D. nebulosus Cdz., type unique [Cand.};
Dilobitarsus nebulosus Cand., type ex. coll. Castelnau [Gahan] (BMNH).

Dilobitarsus nubilus Candéze
Dilobitarsus nubilus Candéze, 1857 : 83.
LECTOTYPE (present designation). CoLtompBia: g, Laf. N. Grenada; Janson

coll. 1903 : 130; Nov. Grenada, Goudot; Dilobitarsus nubilus Cdze., Cand., type
e coll. de Laferté [Janson] (BMNH).

Paralectotype. 9, Laf. N. Grenada; Janson coll. 1903: 130; Dilobitarsus
nubilus Cdze, Cand., type e coll. de Laferté [Janson] (BMNH).

Dilobitarsus pendleburyi Fleutiaux
Dilobitarsus pendleburyi Fleutiaux, 1934d : 178.
Holotype: g, N. Borneo, Bettotan, Nr. Sandakan, Aug. 13, 1927/C.B.K. &
H.M.P., F.M.S. Museums; Brit. Mus. 1934.80 [=presented by Imp. Inst. Ent.];

Dilobitarsus pendleburyi Fleut., type [Fleut.] (BMNH). N. Grenada = Colombia,
see D. colombianus, p. 96.

Dilobitarsus petiginosus Germar
Dilobitarsus petiginosus Germar, 1840 : 246; pl. 1, fig. 3.
The description is based on an unrecorded number of specimens from Brazil.

Type-material: not located. For the history of the Germar collection see p. 000.
The generic attribution is based on the figure.

Dilobitarsus quadrituberculatus Candéze

Dilobitarsus quadrituberculatus Candéze, 1857 : 81.

The description is based on an unrecorded number of specimens from St Catherine,
Brasil.

Type-material: not found in BMNH. ?IRSNB, Brussels.

Generic attribution is based on the following specimen: 3, Laf. Minas Ger. Brasil;
Janson coll. 1903 : 130; Dilobitarsus quadrituberculatus Cdze, Cand., e coll. de
Laferté [Janson] (BMNH). This specimen agrees well with Candéze’s description
of the variety with longitudinal red marks on the elytra and may be the specimen
on which he based the description.

100 CPE VE VON TERAWYAB Ks

Dilobitarsus subsulcatus Candéze
Dilobitarsus subsulcatus Candéze, 1874 : 34.

LECTOTYPE (present designation). Ecuapor: 3g, Ecuador/Macas. [Janson];
Janson coll. 1903.130; D. subsulcatus Cdz., type [Cand.]; Dilobitarsus subsulcatus
Cand. type. Ecuador [Gahan] (BMNH).

The published locality is Equateur, the French form of Ecuador. Janson
obviously re-labelled the specimen to conform with the accepted English rame of
the locality.

Dilobitarsus sulcicollis (Solier) comb. n.
Anancantha sulcicollis Solier, 1851 : 18.

LECTOTYPE (present designation). CHILE: 3, Museum Paris, Chili, Cl. Gay,
1845; Anacantha sulcicollis Sol. Chili. [Solier]; type du Solier [Fleut.] (MNHN,
Paris).

Fleutiaux (1907 : 169) states that he has no hesitation in synonymizing A delocera
angustata F. Philippi, 1861 with swlcicollis Solier despite the fact that he has not
seen the type of the former (? NM, Santiago, Horn & Kahle, 1936: 208). I have
been unable to obtain a copy of the original description of angustata. It appears
to have been based on material collected at Valdivia (F. Philippi, 1887 : 83).

Dilobitarsus tessellatus Candéze
Dilobitarsus tessellatus Candéze, 1874 : 32.

LECTOTYPE (present designation). Brazit: 9, N. Frib.; Janson coll.
1903 . 130; Dilobitarsus tessellatus Cdz. [Cand.]; Dilobitarsus tessellatus Cand.,
type. N. Fribourg [Gahan]; Dilobitarsus tessellatus Cdze in litt. ined. coll.
Deyrolle N. Fribourg [Janson] (BMNH).

Dilobitarsus vitticollis (Fairmaire & Germain) comb. n.

Adelocera vitticollis Fairmaire & Germain, 1860 : 268.
Anacantha vitticollis (Fairmaire & Germain) Candéze, 1874 : 37.
Lacon (Cornilacon) vitticollis (Fairmaire & Germain) Golbach, 1969 : 158.

The description is based on an unrecorded number of specimens from Conception
[CHILE]. Fleutiaux (1907): 167) records that ‘le type primitif de Fairmaire’
was acquired by the IRSNB, Brussels, with Candéze’s last collection. This has
not been confirmed.

The generic attribution is based on the following specimen: 3, Chili coll. Germ.;
Janson coll. 1903 : 130; Adelocera vitticollis L.f. [unknown mss.] Rev. & Mag.
Zool. 1840 [sic], p. 268 [another mss.] (BMNH).

The specimen agrees well with the description and is not conspecific with
D. crux (Philippi) (q.v.).

——

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 101

ELASMOSOMUS Schwarz

Elasmosomus Schwarz, 1902b: 212. Type-species: Elasmosomus fasciculatus Schwarz, by
subsequent designation (Hyslop, 1921 : 643).

GENERIC DIAGNOSIS. Each claw with a group of setae near the base (Text-fig. 11). Tibial
spurs absent (Text-fig. 10). Mesepisternum does not form part of the margin of the mesocoxal
cavity; mesepimeron forms part of the margin of the mesocoxal cavity (Text-fig. 3). Second
and third antennal segments not subequal, the third resembles the fourth and following
segments. Antennal groove shallow and short, not extending beyond the anterior half of the
prosternopleural suture. Body clothed with narrow scales. Prothorax cylindrical, elongate,
not constricted behind the anterior angles. Lateral carina present, pronotal tubercles present
or absent. Propleurae and metasternum without well defined grooves for the reception of the
anterior and middle tarsi. Scutellum simple, without longitudinal carinae. Third and fourth
tarsal segments with ventral lobes.

RANGE OF VARIATION FOUND WITHIN THE GENUS. All the species known to
me bear a strong resemblance to one another. Examination of long series of
specimens suggests that there may be a wide range of variation in the length of
individual specimens belonging to the same species.

HISTORY OF THE GENUS. The genus was established for the African species
attributed to the genus Dzlobitarsus, which originally included only South American
species.

Schwarz originally included the genus in the tribe Dicrepidini but four years
later (1906 : 15) he transferred it to the Agrypnini [sic, see p. 4] without com-
ment. Fleutiaux (1941), who on a previous occasion (1919 : I1) did not recognize
Elasmosomus as a valid genus, preferring to use Dilobitarsus for both African and
South American species, treated them as two entirely distinct genera. He included
Dilobitarsus in the Agrypnitae [sic, see p. 4] but excluded Elasmosomus, basing
his action on the fact that the structure of the head of Elasmosomus species bears
a certain resemblance to that found in Anaspasis (S. America) and Protelater
(New Zealand). Examination of the type-species of these two genera has shown
that the resemblance is only superficial. Both genera differ from Elasmosomus
in that they possess tibial spurs and lack setae at the base of the claws. Further
studies may show that Elasmosomus should not be included in the Agrypninae
but at the present time there is, in my opinion, no convincing reason for excluding
the genus from the subfamily.

The main difference between Elasmosomus and Dilobitarsus, apart from the
distribution, is that in the former the antennal grooves are distinctly shorter
and shallower than in the latter. Only the characteristic cylindrical shape of
the body, the tubercles on the head and the fact that the frons is not margined
anteriorly distinguishes Elasmosomus from Hemicleus. However as no inter-
mediate forms are known and there is no difficulty in assigning species to the
appropriate genus, the two genera have been retained in the present work.

DistripuTion. Africa and Madagascar.

BIoLoGy AND HABITS. Nothing has been published on the life history and
habits of Elasmosomus species. A few specimens known to me bear labels indicating
that they were found on trees or in tree stumps.

|

102 C. M. F. von HAYEK

SEXUAL DIMORPHISM. The species known to me do not appear to display any
distinct sexual dimorphism.

SPECIES INCLUDED IN THE GENUS

The Schenkling catalogue (1925 : 18) lists the names of twenty-six species, of
which three are regarded as synonyms. Three species have been described since
the publication of the catalogue. The generic attribution of all these species
has been confirmed, either by examination of the type-material, determined
material or from the description. It has not been possible to confirm all the
synonymies. One species, Dilobitarsus fleutiauxi Paulian, 1947, is here transferred
to Elasmosomus. As it is a secondary junior homonym of E. fleutiauxi Schwarz,
1906, the replacement name delamarei is proposed.

Elasmosomus alluaudi (Candéze)

Dilobitarsus alluaudi Candéze, 1895p : 52.
Elasmosomus alluaudi (Candéze) Schwarz, 19026 : 297.

The description is based on an unrecorded number of specimens from Madagascar,
Diego-Suarez, Montagne d’Ambre collected and submitted by Alluaud.

Syntypes examined: MADAGASCAR: I 9, Madagascar, Diego Suarez 17, Ch.
Alluaud, 1893; Dilobitarsus alluaudi Cand. [Cand.] Cand. det. [Fleut.]. 1 9, same
locality and collector; alluaudi Cand. Comparé au type, Mus. Brux. 1902 [Fleut.];
Dilobitarsus alluaudi Cand., ex type. 5 ex., same locality and collector (MNHN,
Paris).

The designation of a lectotype is postponed until the ‘type’ material in the
IRSNB, Brussels, to which Fleutiaux’s label and Fleutiaux (1902c; 298) refers,
has been examined.

Elasmosomus apicalis Schwarz

Elasmosomus apicalis Schwarz 1905b : 275.

The description is based on an unrecorded number of specimens from [TANZANIA]
Usambara, Nguela.

Type-material: ?>DEI, Eberswalde.
The generic attribution is based on the description.

Elasmosomus bacillus (Candéze)

Dilobitarus bacillus Candéze, 1881 : 3.
Elasmosomus bacillus (Candéze) Schwarz, 1902b : 214.

The description is based on an unrecorded number of specimens from ABYSSINIA,
collected by Raffray.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 103

Type-material: IRSNB, Brussels, according to Fleutiaux (1902c : 298). Not
confirmed.

The generic attribution is based on the description.

Elasmosomus bituberculatus (Fleutiaux)

Dilobitarsus bituberculatus Fleutiaux, 1902c : 299.
Elasmosomus bituberculus Schwarz, 1906: 15. [Unjustified emendation.]}

LECTOTYPE (present designation). SAN THoME: I 9, San Thomé, Dilo-
bitarsus, bituberculatus Fleut., type [Fleut.] (MNHN, Paris).

Elasmosomus brunneiventris Schwarz
Elasmosomus brunneiventris Schwarz, 1902b : 217.

The description is based on an unrecorded number of specimens from Kamerun
[CAMEROUN].

Type-material: ?>DEI, Eberwalde.

The generic attribution is based on the following specimen: 1 9, Kamerun,
Conradt; brunneiventris Schw. det. Quelle [?Quelle] (MNHN, Paris).

Elasmosomus cornutus (Candéze)

Dilobitarsus cornutus Candéze, 1865 : 8.
Elasmosomus cornutus (Candéze) Schwarz, 1902b : 314.

Candéze recorded the locality as Guinée (probably in the old commercial sense;
the name was commonly used to denote the Atlantic coastal region of Africa from
Cape Verga, 10°18’N. to Cape Negro, 15°45’S.). He also refers to a specimen
from Old Calabar submitted by Murray and another, presumably from the same
locality, in the Saunders collection.

Type-material: IRSNB, Brussels, according to Fleutiaux (1902c : 298). Not
confirmed.

According to Horn & Kahle (1936: 186, 239, 380) Murray’s and part of
Saunders’ collections are in the BMNH. The BMNH does not possess any
cornutus specimens from these collections or from the Candéze collection. The
remainder of the Saunders collection is in the Fleutiaux collection (MNHN, Paris).
Not located, 1971.

The generic attribution is based on the following specimen: ConGo (BRAZZAVILLE):
I 9, Benito, Congo Franc.; cornutus Cand. comparé type, Mus. Brux. 1902 [Fleut.]

(MNHN, Paris).
Elasmosomus cylindricus (Fleutiaux)

Dilobitarsus cylindricus Fleutiaux, 1902c : 300.
Elasmosomus cylindricus (Fleutiaux) Schwarz, 1906 : 15.

104 ; CIM. Eo von BELA YE Ke

LECTOTYPE (present designation). CONGO (BRAZZAVILLE): 9, Benito, Congo
Franc; Dilobitarsus cylindricus Fleut., type (Fleut.) (MNHN, Paris).

Elasmosomus delamarei nom. n.

Diolobitarsus fleutiauxi Paulian, 1947 : 7, fig.
Elasmosomus fleutiauxit (Paulian) comb. n. [Secondary junior homonyn of Elasmosomus
fleutiauxt Schwarz, 1906. ]

Holotype. Ivory Coast: 9, Céte d'Ivoire, Reserve du Blanco, R. Paulian &
G. Delamare; Reseu nogq. fre. 8": [illegible]; Dilobitarsus [Fleut.]; Dilobitarsus
fleutiauxi n.sp. [Paulian] (MNHN, Paris).

The published locality is: Cote d'Ivoire, reserve forestiére du Banco en battent
la lisiere d’une clariere 4 8 heurs du matin, ix.1945, R. Paulian et C. Delamare-
Debouttville.

Elasmosomus edmundi Quelle

Elasmosomus edmundi Quelle, 1955 : 228.

Holotype. CAMEROUN: Kamerun, Johann Albrechtshédhe (Conradt, 1897)
(NMHU, Berlin). Not examined.

The generic attribution is based on the description.

Elasmosomus fasciculatus Schwarz

Elasmosomus fasciculatus Schwarz, 1902b : 214.

The description is based on an unrecorded number of specimens from Kamerun
[CAMEROUN].

Type-material: ?>DEI, Eberswalde.

The generic attribution is based on the description.

Elasmosomus filiformis (Candéze)

Dilobitarsus filuformis Candéze, 1882 : 3.
Elasmosomus filiformis (Candéze) Schwarz, 19026 : 214.

The description is based on an unrecorded number of specimens from ABYSSINIA
collected by Raffray.

Type-material: IRSNB, Brussels according to Fleutiaux, r1902c: 299. Not
confirmed.

The generic attribution is based on the description.

Fleutiaux (1902c : 299) regards this species as a synonym of E. vaffrayi (Candéze,
1878). The synonymy has not been confirmed.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 105

Elasmosomus filum (Candéze)

Dilobitarsus filum Candéze, 1897 : 7.
Elasmosomus filum (Candéze) Schwarz, 1902b : 214.

The description is based on an unrecorded number of specimens from GABON.

Type-material: IRSNB, Brussels, according to Fleutiaux (1g02c : 298). Not
confirmed.

The generic attribution is based on the description.

Elasmosomus fleutiauxi Schwarz

Dilobitarsus linearis Fleutiaux, 1902c : 301.
Elasmosomus fleutiauxi Schwarz, 1906: 15. [Replacement name for Elasmosomus linearis
(Fleutiaux), secondary junior homonym of E. linearis Schwarz 1902b : 219.]}

LECTOTYPE (present designation). COoNGo (BRAZZAVILLE): 2, Benito, Congo
Franc; Dilobitarsus linearis Fleut, type [Fleut] (MNHN, Paris)

Elasmosomus humeralis Schwarz

Elasmosomus humeralis Schwarz, 1902b : 218.

The description is based on an unrecorded number of specimens from Kamerun
[CAMEROUN]

Type-material: ? DEI, Eberswalde
The generic attribution is based on the description.

Elasmosomus jeanneli Mouchet
Elasmosomus jeanneli Mouchet, 1949 : 113.
Holotype. Mapacascar: 2, Madagascar. Coll. A. Sicard, 1930; Mont d’Ambre;
Décembre; Elasmosomus jeanneli, type [Mouchet] (MNHN, Paris).

Paratypes examined: II specimens bearing ‘cotype’ labels: 1 ex., same date,
locality and collection as holotype. I ex., same date and locality as holotype.
3 ex., same locality and collection as holotype, one specimen bears the date ‘octobre’.
I ex., Février; Madagascar Coll. A. Sicard, 1930. 5 ex., same collection as holotype,
four specimens are labelled ‘Dilobitarsus’ (MNHN, Paris).

The Mouchet collection also contains paratypes (see description). Mouchet does
not record the number of paratypes in each collection.

Elasmosomus linearis Schwarz

Elasmosomus linearis Schwarz, 1902) : 218.

The description is based on an unrecorded number of specimens from Kamerun
[CAMEROUN]

Type-material: ? DEI, Eberswalde.
The generic attribution is based on the description.

106 Cc. M. EF. von HAYEK

Elasmosomus mirificus (Candéze)

Dilobitarsus mivificus Candéze, 1895 : 48.
Elasmosomus mivificus (Candéze) Schwarz, 1902b : 214. .

The description is based on a single specimen from ToGo submitted by Kraatz.
Holotype: IRSNB, Brussels according to Fleutiaux (1902c : 298). Not confirmed.
The generic attribution is based on the description.

Elasmosomus mocquerysi (Fleutiaux)

Dilobitarsus mocquerysi Fleutiaux, 1902¢ : 300. |
Elasmosomus mocquerysi (Fleutiaux) Schwarz, 1906 : 15.

LECTOTYPE (present designation). SAo THOME: 9, Sao Thomé; Dilobitarsus
mocquerysi Fleut., type [Fleut.]; Ann. Soc. Ent. Belge 1902 pp. 297 et 300 [Fleut.]
(MNHN, Paris).

Paralectotype: I ex., Sao Thomé (MNHN, Paris).

Elasmosomus parallelus Schwarz
Elasmosomus parallelus Schwarz, 1902b : 218.

The description is based on an unrecorded number of specimens from Kamerun
[CAMEROUN].

Type-material: ? DEI, Eberswalde.
The generic attribution is based on the description.

Elasmosomus pauliani Mouchet
Elasmosomus pauliani Mouchet, 1949 : 113.

Holotype. Mapacascar: Baie d’Antongil, M. Mocquerys, 1898; Elasmosomus
pauliani, type [Mouchet] (MNHN, Paris).

Paratype: 9, same locality as the holotype (Mouchet collection). Not examined.

Elasmosomus pulchellus Schwarz
Elasmosomus pulchellus Schwarz, 19050 : 273.

The description is based on an unrecorded number of specimens from Toco.
Type-material: ? DEI, Eberswalde.
The generic attribution is based on the description.

Elasmosomus pulcher Schwarz

Elasmosomus pulcher Schwarz, 1905p : 275.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 107

The description is based on an unrecorded number of specimens from Kamerun,
Balanga [CAMEROUN].

Type-material: ? DEI, Eberswalde.
The generic attribution is based on the description.

Elasmosomus raffrayi (Candéze)

Dilobitarsus vaffrayi Candéze, 1878) : LII (6).
Elasmosomus raffrayi (Candéze) Schwarz, 19026 : 214.

The description is based on two specimens from Zanzibar [TANZANIA] collected
by Raffray.

Syntypes: 1 ex., Candéze collection (IRSNB, Brussels). Not examined. I ex.,
MCZN, Genoa.

The generic attribution is based on the following specimen: 14, Zanzibar, Raffray;
Dil. raffrayi Cand. [Cand.]; Coll. Alluaud, ex type raffrayi Cand. [Fleut.] (MNHN,
Paris). Unless one of the two syntypes believed to be in the IRSNB, Brussels and
MCZN, Genoa are found to be missing, this specimen cannot be part of the syntype
series.

Fleutiaux (1902c : 299) lists E. ramusculus (Candéze) described from Togo (see
below) and E. filiformis (Candéze) from Abyssinia (see p. 104) as synonyms. The
synonymies have not been confirmed.

Elasmosomus ramusculus (Candéze)

Dilobitarsus ramusculus Candéze, 18954 : 47.
Elasmosomus vamusculus (Candéze) Schwarz, 1902b : 214.

The description is based on a single specimen from Toco submitted by Kraatz.

Holotype: IRSNB, Brussels according to Fleutiaux, Igo2c : 299. Not confirmed.
Fleutiaux (1902c : 299) places this species in synonymy with raffrayi Candéze.
The synonymy has not been confirmed.

Elasmosomus signifer (Candéze)
Dilobitarsus signifer Candéze, 18956 : 53.

Dilobitarsus perrieri Fairmaire, 1901 : 169. [Synonymized by Fleutiaux, 1902c : 299.]
Elasmosomus signifer (Candéze) Schwarz, 1902b : 214.

Dilobitarsus signifer Candéze. Holotype. Mapacascar: 9, Madagascar, coll.
Ch. Alluaud; D. signifer Cand. [Cand.] Cand. det. (MNHN, Paris).

The published locality is Diego-Surarez, Montague d’Ambre.

Dilobitarsus perrieri Fairmaire. LECTOTYPE (present designation), Mapa-
GASCAR: Perrier [Fairm.]; Museum Paris, Madagascar, Perrier de la Bathie;
Dilobitarsus [Fleut.] perrieri Frm., type [Fairm.]; signifer Cand., minor, perrieri
Fairm. [Fleut.] (MNHN, Paris).

The published locality is Madagascar, Bélumbé (H. Perrier).

108 Cl. Ey “Vion eH AYE

Elasmosomus sobrinus (Candéze)

Dilobitarsus sobvinus Candéze, 18954 : 45.
Elasmosomus sobrinus (Candéze) Schwarz, 1902b : 214.

Holotype: Toco, submitted by Kraatz. IRSNB, Brussels according to
Fleutiaux. (1902c: 298). Not confirmed.

The generic attribution is based on the following specimen: 9. Togo. D. sobrinus
Cand. [Cand.]; Coll. Alluaud, ex type Cand. [Fleut.] (MNHN, Paris). This specimen
cannot be part of the type-series as Candéze records only a single specimen.

Elasmosomus undulatus Schwarz
Elasmosomus undulatus Schwarz, 1902b : 215.

The description is based on an unrecorded number of specimens from Kamerun
[CAMEROUN].

Type-material: ? DEI, Eberswalde.
The generic attribution is based on the description.

Elasmosomus variegatus (Fleutiaux)

Dilobitarsus variegatus Fleutiaux, 1902c : 301.
Elasmosomus variegatus (Fleutiaux) Schwarz, 1906 : 15.

LECTOTYPE (present designation). CoNGo (BRAZZAVILLE): 2, Benito, Congo
Franc; Dilobitarsus variegatus Fleut., type [Fleut] (MNHN, Paris).

Elasmosomus vicinus (Candéze)

Dilobitarsus vicinus Candéze, 1895b : 53.
Elasmosomus vicinus (Candéze) Schwarz, 19026 : 214.

Syntypes examined: MADAGASCAR: I ex., Madagascar, Diego-Suarez, 17, Ch.
Alluaud 1893; D. vicinus Cand., Cand. det. [Cand.]. 6 ex., same locality without
determination labels. 1 9, Diego Suarez, Alluaud; vicinus Cand., comparé au type
Mus. Brux. 1902 [Fleut.]; Dilobitarsus vicinus Cand., ex type [Fleut.]. 1 ex.,
same locality without determination label (MNHN, Paris).

The description is based on an unrecorded number of specimens from Diego
Suarez, Montagne d’Ambre, collected and submitted by Alluaud. Fleutiaux
(Ig02c : 298) and the note on one specimen referring to the ‘type’ in the IRSNB,
Brussels, suggests that Candéze did not return all the material to Alluaud, but
retained a specimen or specimens for his own collection. Designation of the lecto-
type is postponed until this material has been examined.

HEMICLEUS Candéze
Hemicleus Candéze, 1857 : 180. Type-species: Hemicleus caffey Candéze, by monotypy.

GENERIC DIAGNOSIS. Each claw with a group of setae near the base (Text-fig. 11). Mesepi-
sternum does not form part of the margin of the mesocoxal cavity; mesepimeron forms part of

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 109

the margin of the mesocoxal cavity (Text-fig. 3). Second and third antennal segments not
subequal, the third segment resembles the fourth. Antennal groove shallow and short, not
extending beyond half the length of the sternopleural suture. Body clothed with narrow
scales. Prothorax moderately convex but not cylindrical; without constriction behind the
anterior angles. Lateral carinae present. Pronotum without tubercles. Propleurae and
metasternum without well defined grooves for the reception of the anterior and middle tarsi.
Scutellum simple, without carina. Third and fourth tarsal segments with ventral lobes.

Structurally the genus bears a certain resemblance to Elasmosomus and is distinguished from
it only by its general appearance, the body being less cylindrical and the pronotum entirely
devoid of tubercles. Within the subfamily these characteristics are not generally of any value
at the generic level, but as there is no difficulty in assigning the known species to one genus or
the other both genera have been retained in the present work.

HISTORY OF THE GENUS. The genus was established for a single species. It
has gained general acceptance and over the years a number of new species have
been described.

SEXUAL DIMORPHISM. The species appear to be indistinguishable externally.
DISTRIBUTION. Africa.

BIOLOGY AND HABITS. Nothing is known of the life history and habits of
Hemuicleus species.

SPECIES INCLUDED IN THE GENUS

Schenkling (1925 : 20 & 514) lists thirteen species and six have been described
subsequently. The generic attribution of sixteen species has been confirmed.
One species, planus Fleutiaux is transferred to Adelocera and another, jeanneli
Fleutiaux transferred from the same genus. The remaining two species (adspersulus
Klug and majusculus Candéze) are unknown to me.

Hemicleus apicalis Schwarz

Hemicleus apicalis Schwarz, 1898b : 183

Syntype examined. TANZANIA: 9, Usamb. Weise; Coll Schwarz; Typus; apicalis
Schw. [Schwarz] (DEI, Eberswalde).
The published locality is Usambara.

Hemicleus caffer Candéze
Hemicleus caffery Candéze, 1857 : 181; pl. 3, fig. 3.
The description is based on an unrecorded number of specimens from ‘Cafrerie’
[Sout AFrricA] submitted by Dohrn.

Type-material. Not located in BMNH. ? IRSNB, Brussels (see p. 271) or
) possibly in the Dohrn collection in the IZPAN, Warsaw.

The generic attribution is based on the following specimen; 1 4, illegible locality;
Afr. Sud: Hemicleus caffer Cand. [Fleut.] (MNHN, Paris).

110 Cc. M. F. von HAYEK

Hemicleus cardiophoroides Fleutiaux
Hemicleus cardiophoroides Fleutiaux, 1919 : 13.

LECTOTYPE (present designation). KrNyaA: 9, Afr. or. Anglaise, Taveta,
Alluaud & Jeannel, Mars 1912, st. 65; Hemicleus cardiophoroides Fleut., type
[Fleut.] (MNHN, Paris).

Paralectotype. TANZANIA: 9, Afr. or. Allemande; Kilimandjaro, versant sud-est,
Alluaud & Jeannel; Zone inferieure, Neu Moschi, 800 m. Avril Ig12. st. 72;
conforme a caffer in coll. Cand. Afr. orient. J. Hub... . [illegible]; Nov. 1930;
non le type [Fleut.] (MNHN, Paris).

The meaning of Fleutiaux’s note on the paralectotype is not clear. There does
not seem to be any obvious difference between this specimen and the lectotype.

Hemicleus chappuisi Fleutiaux
Hemicleus chappuisi Fleutiaux, 1935) : 197: fig. 3.

LECTOTYPE (present designation). ETHIOPIA: 9, Ethiopie Merid. Nanaropus,
Bords du Rudolphe, 565m; Museum Paris, Mission de L’Omo. C. Arambourg,
P.-A Chappuis & R. Jeannel; Hemicleus chappuisi Fleut., type [Fleut.] (MNHN,
Paris).

Hemicleus dorsalis Schwarz
Hemicleus dorsalis Schwarz, 18984 : 183.

Syntypes examined. CAMEROUN: I ex., Kameroun; Coll. Schwarz; Typus;
dorsalis Schw. [Schwarz] (DEI, Eberswalde). I ex., Kamerun Conrad;
Hemicleus dorsalis Schw. [Fleut.] (MNHN, Paris). I have no doubt that this
specimen originally formed part of the syntype-series and that it was subsequently
acquired by Fleutiaux.

The description is based on an unrecorded number of specimens collected by
Conrad and submitted by Kraatz.

Hemicleus duplicatus nom. n.

Hemicleus jeannelt Fleutiaux, 1935) : 196; fig. 2. [Junior secondary homonym of Hemicleus
jeanneli (Fleutiaux, 1919).]

LECTOTYPE (present designation). KEnyaA: 9, Kenya, Env. de Lokitanyalla,
West Suk-Turkana, 1,200m; Hemicleus duplicatus Fleut., n.n.p. jeanneli Miss.
Omo 1935 [Fleut.] (MNHN, Paris).

Fleutiaux appears to have been aware that Adelocera jeanneli Fleutiaux, 1919
should be assigned to Hemicleus (see p. 111) and that H. jeanneli (Fleutiaux, 1935)
therefore required a new name. Although he altered the names on the specimens
he did not publish the new combination or the new name.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 111

Hemicleus elegans Fleutiaux

Hemicleus elegans Fleutiaux, 1935c : 94.

LECTOTYPE (present designation). ? E. Arrica: 9, R. E. Dent, Chania R.
7800, July 1929; Hemicleus elegans Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 1 3, R. E. Dent, Chania R. 7800, July 1929; Hemicleus elegans
Fleut., co-type [Fleut.] (MNHN, Paris). 1 g, R. E. Dent, Ghania [sic] R. 7,800,
July 1929. Hemicleus elegans Fleut. [Fleut.| (NM, Nairobi).

The position of the published locality, Chania River, is unknown to me.

Hemicleus fasciculatus Schwarz

Hemicleus fasciculatus Schwarz, 19050 : 276.

Syntype examined. CAMEROUN: 9, Kamerun, Barombi, Conrad; Coll. Schwarz;
Typus; fasciculatus Schw. [Schwarz] (DEI, Eberswalde).

Hemicleus ferrantei Buysson

Hemicleus fevvantet Buysson, 1911 : 19.

The description is based on an unrecorded number of specimens from [EGYPT]
‘Luxor, premiers jours de juillet 1908, 1909, 1910 (coll. Ferranti).’ The Ferranti
collection is now in the possession of the Entomological Society of Egypt (Horn &
Kahle, 1935: 74). Not confirmed.

Syntypes examined. 1 4g, Luxor, 2.vii.10; Luxor, Ferranti; Hemicleus ferrantei
Du B. Egypte [du Buysson]. 2 g, Luxor, 7.vii.og, G. Ferranti (MNHN, Paris).

These specimens were presumably retained by du Buysson and were acquired by
the MNHN, Paris with his collection.

The designation of the lectotype is postponed until the remainder of the series
has been examined.

Hemicleus inordinatus Fleutiaux
Hemicleus inordinatus Fleutiaux, 19320 : 2.
LECTOTYPE (present designation). MozamBiguE: g, Museum Paris, Afr.

or. port., Nova Choupanga, Dr Lesne, 1928; Avril; Boite 16; Hemicleus inordinatus
Fleut., type [Fleut.] (MNHN, Paris).

Hemicleus jeanneli (Fleutiaux) comb. n.
: Adeloceva jeanneli Fleutiaux, 1919 : Io.
: Holotype. Tanzania: 9, Afr. or. Allemande, Kilimandjaro, versant sud-est,
_ Alluaud & Jeannel; Zone inferieur, Neu Moschi, 800 m, Avril 1912, st. 72; Adelocera

jeanneli Fleut., type [Fleut.]; Hemicleus non Adelocera [Fleut.] (MNHN, Paris).
Fleutiaux did not publish the transference of this species to Hemicleus.

II2 GC. M. Bo von EPAVEK

Hemicleus longicollis Schwarz
Hemicleus longicollis Schwarz, 1905b : 277.
Syntype examined. ETHIOPIA: J, Galla, R. V. Erlanger; Coll Schwarz; Typus;
longicollis Schw. [Schwarz] (DEI, Eberswalde).

The published locality is Somali. It seems probable that the name Galla refers
to the tribe inhabiting the country 150 miles south of Addis Ababa and not the
hills of the same name 20 miles south of the equator, 40°E., in Kenya.

Hemicleus normandi Buysson
Hemicleus normandi Buysson, 1920 : 12.
LECTOTYPE (present designation). Tunisia: T. le Kef, Dr Normand; Hemi-
cleus normandi Buyss [Buyss.| (MNHN, Paris).
Paralectotype: 1 3, T. le Kef, Dr Normand, Hemicleus normandi H. du Buyss;
co-type (MNHN, Paris).
The published data are: Tunisia, le Kef sur les herbes séches, en aout (Dr Normand.)

Hemicleus ordinatus Fleutiaux
Hemicleus ordinatus Fleutiaux, 19320 : 2.
LECTOTYPE (present designation). MozAMBIQUE: gj, Museum Paris, Zambeze,

Nova Choupanga, Pres Chemba, P. Lesne; Janvier; Hemicleus ordinatus Fleut.,
type [Fleut.] (MNHN, Paris).

Hemicleus quadricollis Fleutiaux
Hemicleus quadricollis Fleutiaux, 191g : 12.

Holotype. Kenya: 9, Afr. or. Ang. (Wa Kikuyu) Fort Hall, Alluaud & Jeannel,
Janv. 1919, 1330 mst. 30; Hemicleus quadricollis Fleut., type [Fleut.] (MNHN, Paris).

Hemicleus villiersi Cobos
Hemicleus villierst Cobos, 1970 : 134; figs 1, 2.
Holotype. CoNGo (BRAZZAVILLE): 3, Congo, Sibiti, xi [sic] 1963; Museum

Paris, Mission A. Descarpentries et A. Villiers, 1963-64; Hemicleus villiersi N. sp.
Holotype [Cobos] (MNHN, Paris).

Paratype. I 4, Sibiti, xi1.63 (Cobos coll. Almeria, Spain). Not examined.
The published date is xi1.63.

Hemicleus weisei Schwarz
Hemuicleus weiset Schwarz, 1898b : 183.

Syntype examined. TANZANIA: sex undetermined, Usamb. Weise; Coll Schwarz;
Typus; weisei Schw. [Schwarz] (DEI, Eberswalde).
The published locality is Usambara.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 113

AGRYPNUS Eschscholtz

Agrypnus Eschscholtz, 1829 : 32. Type-species: Elater murvivus Linnaeus, by subsequent
designation (Westwood, 1838: Synopsis, 26).

[Lacon sensu Germar, 1840 and subsequent authors, not Castelnau, 1836. Misinterpretation.]

Mecynocanthus Hope, 1837 : 53. Type-species: Mecynocanthus unicoloy Hope, by monotypy.
Syn. n.

Tylotarsus Germar, 1840: 247. Type-species: Tylotarsus cinctipes Germar, by monotypy.
Syn. n.

Tilotarsus Candéze, 1857 : 170. [Unjustified emendation. |

Myrmodes Candéze, 1857 : 168. Type-species: Myrmodes akidifoymis Candéze, by monotypy.
Syn. n.

Ayrchontas Goezis, 1886 : 23. Type-species: Elater murinus Linnaeus, by monotypy.

Pseudolacon Blackburn, 1890 : 89. Type-species: Pseudolacon rufus Blackburn, by monotypy.
Syn. n.

Homeolacon Blackburn, 1890 : 90. Type-species: Homeolacon gracilis Blackburn, by monotypy.
Syn. n.

Lobotarsus Schwarz, 1898a : 131. Type-species: Lobotarsus decovatus Schwarz, by subsequent
designation (Hyslop, 1921 : 263). Syn. n.

Lobitarsus Fleutiaux, 1935¢c : 93. [As asubgenus of Tylotarvsus.| [Unjustified emendation. ]

Enoploderes Schwarz, 1898a : 131. Type-species: Elater (Conoderus) cuspidatus Klug, by
monotypy. [Junior homonym of Enoploderes Faldermann, 1837.]

Centrostethus Schwarz, 1898a : 414. [Replacement name for Enoploderes Schwarz.] Syn. n.

Compsolacon Reitter, 1905: 6. Type-species: Elater crenicollis Ménétriés, by monotypy.
{[Synonymized with Lacon auct. by Fleutiaux, 1918d : 198.]

Paralacon Reitter, 1905: 6. Type-species: Lacon cinnamomeus Candéze, by monotypy.
[Reduced to subgenus of Agrypnus Eschscholtz, Ohira, 1968) : 364.) Syn. n.

Neolacon Miwa, 1929 : 235. Type-species: Neolacon formosanus Miwa, bymonotypy. [Reduced
to subgenus of Compsolacon Reitter, Miwa, 1934: 14.) Syn. n.

Colaulon Arnett, 1952 : 116. Type-species: Elater rectangularis Say, 1925, by original designa-
tion. Syn. n.

Cryptolacon Nakane & Kishii, 1955: 1. Type-species: Cryptolacon myamoti Nakane & Kishii,
1955, by original designation. [Reduced to subgenus of Colaulon by Kishii, 1961 : 25.]
Syn. n.

Sabikikorius Nakane & Kishii, 1955 : 3. [as subgenus of Agrypnus Eschscholtz]. Type-species:
Lacon fuliginosus Candéze, 1865 : 10, by original designation. [Raised to generic status,
Kishii, 1964 : 15.) Syn.n.

Sagojyo Kishii, 1964 : 30 [as a subgenus of Colaulon Arnett]. Type-species: Colaulon (Sagojyo)
yuppe Kishii, by original designation. [Raised to generic status, Ohira, 1967) : 103.]
Syn. n.

Sagojo Ohira, 1968b : 364. [Unjustified emendation. }

Archontoides Cobos, 1966 : 651. Type-species: Archontoides pretoriensis Cobos by monoytpy.
Syn. n.

Pyrganus Golbach, 1968: 198. Type-species: Lacon tuspanensis Candéze by original designa-

' tion. Syn. n.

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae near the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepisternum and mesepimeron do not form part of the
| margin of the mesocoxal cavity (Text-fig. 2). Second and third antennal segments small,
subequal, each smaller than the fourth and following segments (Text-fig. 13) Antennal groove
not extending beyond the anterior two-thirds of the prosternopleural suture; depth of groove
variable. Vestiture scale-like. at least in part. Prothorax not constricted behind the anterior
angles: lateral margin of prothorax carinate. Propleural grooves or depressions for the
reception of the tarsi, if present, do not run parallel with the antennal grooves (Text-fig. 7).

H

114 CaM. ©. von) HAYEK

Metasternal grooves or depressions for the reception of the tarsi, if present, directed diagonally
across the metasternum so that the distal ends, if produced, would cut the lateral margin of the
metasternum at a point within the posterior quarter of its length or run directly into the
posterior lateral angles of the metasternum (Text-fig. 5). Scutellum variable in shape, but
never with a longitudinal carina. Tarsi with or without ventral lobes (Text-figs 15, 16).

RANGE OF VARIATION WITHIN THE GENUS. Species possessing this combination
of characteristics display a very wide range of variation in such features as the
size and shape of the fourth to the eleventh antennal segments, the shape and
appearance of the surface and margin of the head, prothorax and elytra, the shape
of the mesosternal groove, the degree of development of the propleural and
metasternal depressions or grooves for the accommodation of the anterior and
middle tarsi, the length of the femora in relation to the width of the body, the
structure of the tarsi and the colour and distribution of the scales clothing the body.

Up to the present time these characteristics have been used, either singly or in
combination with one another, as the diagnostic features of genera and subgenera.
The practice of studying restricted faunas has obscured the fact that, when
considered on a world-wide basis, these characteristics are very variable and do not
appear in any constant relationship to one another.

HISTORY OF THE GENUS. Eschscholtz listed the names of twelve species when he
established the genus. Two species, cribrosus and granulosus were described as new
and two others, senegalensis Dejean and coenosus Hope were undescribed. The
description of coenosus Hope was not published until 1831 and senegalensis Dejean
was eventually listed as a synonym of notodonta Latreille by Candéze (1857 : 27).

Latreille (1834 : 143) included six of Eschscholtz’s Agrypnus species in his inter-
pretation of the genus, which he divided into two parts. The first included species
with a tubercle at the base of the prothorax (E£. fuscipes Fabricius and E. senegalen-
sts Dejean) and the second the species without a tubercle (E. atomarius, varius,
fasciatus and murius). He also included quadrimaculatus Fabricius (= Betarmon
ferrugineus Scopoli) in the second group. The remaining six species included in
Agrypnus by Eschscholtz (tomentosus, luridus, coenosus, conspersus, cribrosus and
granulosus) are not mentioned by Latreille.

Germar (1840 : 253) included four (luridus, tomentosus, fuscipes and atomaria) of
Eschscholtz’s Agrypnus species in his interpretation of the genus and assigned to it
four additional species (notodonta Latreille, ruber Perty, moerens n. sp. (Alaus, see
p- 240) and adspersus Herbst (Aliteus, see p. 240)). Of Eschscholtz’s remaining eight
species he transferred murinus to Lacon and varius, fasciatus and conspersus to
Adelocera. Germar does not refer to coenosus, cribrosus, granulosus or senegalensis.

Candéze (1857) limited the genus to four of the species originally included by
Eschscholtz (tomentosus, fuscipes, luridus and senegalensis) to which he added 26
species bearing a very close resemblance to them.

Candéze’s interpretation of Agrypnus (= Lanelater of the present work, see
p. 240), containing relatively large species with long antennal grooves and clothed
with fine setae, was generally accepted for over sixty years, and Hyslop’s (1921 : 624)
designation of tomentosus Fabricius as the type-species appeared perfectly justified.
It was not until Lane drew Arnett’s (1952 : 105) attention to the fact that Westwood

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE II5

(1838 : 26) had properly designated murinus Linnaeus as the type-species of Agrypnus
Eschscholtz that it became apparent that the correct interpretation of Agrypnus
is entirely different from that of earlier workers.

NOTES ON THE GENERA AND SUBGENERA NEWLY PLACED IN SYNONYMY WITH
Agrypnus.

Mecynocanthus Hope. M. unicolor Hope, the type-species of this monobasic
genus is discussed on p. 225. Apart from the locality (‘India oriental’), which
is probably erroneous, this species does not differ significantly from Centrostethus
cuspidatus (Klug) (see below).

Tylotarsus Germar. The genus was erected for cinctipes, a Madagascan species
with lobed fourth tarsal segments. Schwarz (1898a: 129) redefined the genus,
restricting it to species with lobed fourth tarsal segments in which the margins
of the mesosternal groove are horizontal posteriorly and steeply declivous anteriorly
and the anterior angles of the prothorax produced but not sharply pointed as in
Centrostethus. Examination of the Madagascan material available has shown
that development of the anterior angles of the prothorax varies from one species
to another and bears no constant relationship to the shape of the mesosternal
groove. Schwarz (1903a@ : 359) comments that the shape of the mesosternal groove
of T. angularis Schwarz lies between that found in the type-species of Tylotarsus
and Lobotarsus.

Myrmodes Candéze. The genus is characterized by the sloping humeri of the
elytra. This characteristic is associated with the reduction in length of the wings
and is found, though in a less extreme form, in the Australian Agrypnus octavus
(Candéze) and the South African Agrypnus pretoriensis (Cobos) (= paenulatus
Boheman)). The latter species was used as a basis for the erection of the genus
Archontoides. In each case reduction of the humeri is accompanied by a reduction
in length of the metasternum and the scutellum. Other transitional forms between
Agrypnus and Myrmodes are known. In certain South African (e.g. muscerda
(Candéze)) and Australian (e.g. divaricatus (Candéze)) species, the wings, metasternum
and scutellum are reduced in length, but the humeri are normal.

Pseudolacon Blackburn. The genus was erected for rufus Blackburn which
differs from Lacon sensu auctt. (Agrypnus of this work) in that the body is more
cylindrical and the first four tarsal segments conical. Both characteristics are
too variable to justify the retention of the genus.

Homeolacon Blackburn. The diagnostic features of this genus are the strongly
transverse fourth to eleventh antennal segments and elongate tarsal segments.
The lectotype of gracilis Blackburn, the only specimen known to me, is a male.
It bears a close resemblance to Agrypnus cylindricus (Candéze), (= truncatus
(Herbst)) an Indian species in which the antennal segments of the male are very
much more transverse and the tarsi more elongate, than those of the female. It
seems very probable that the female of gracilis Blackburn, will be found to possess
the moderately serrate antennae found in the majority of Agrypnus species.

I16 Cees, VES vio NG EL ASyoE Is

Lobotarsus Schwarz. The genus was erected for species differing from Yylotarsus
in that the margin of the mesosternal groove is inclined and not angled. Schwarz
(1903a : 359) himself cast doubt on the value of this character when he drew atten-
tion to the fact that the appearance of the margin of the mesosternal groove of
T. angularis Schwarz was intermediate between that found in Tylotarsus and
Lobotarsus. Fleutiaux clearly had misgivings concerning the genus. In 1935¢
he treated Lobitarsus [sic] as a subgenus of Tylotarsus. In 1941 he restored it to
generic status, redefining it on the basis of the shape of, and presence of tubercles
on, the prothorax.

Centrostethus Schwarz was proposed as a new name for Enoploderes Schwarz,
1898, preoccupied by the Cerambycid genus Enoploderes Faldermann, 1837. The
type-species cuspidatus Klug is distinguished from A. murinus (Linnaeus) only by
the lobed tarsi and sharply pointed anterior angles of the prothorax.

Compsolacon Reitter. The genus is characterized by the crenulate lateral margins
of the prothorax. This characteristic is known to vary intraspecifically and also
on the two sides of the same individual. Fleutiaux (1918d : 198) placed the genus
in synonymy with his interpretation of Lacon (= Agrypnus of the present work).
Since that time there has been little agreement concerning the status of Compsolacon.
In the most recent work, that of Van Zwaluwenburg (1966 : 298), Compsolacon
is reduced to a subgenus of his interpretation of Adelocera Latrielle, which corresponds
to Agrypnus Eschscholtz of the present work.

Paralacon Reitter. Reitter erected this genus for his interpretation of Lacon
cinnamomeus Candéze, which was probably correct. In this species the posterior
femora are elongate, so that the femoro-tibial articulations are visible from above
when the beetle is at rest. This characteristic is developed to a greater or less
degree in a number of species which in all other respects bear a close resemblance
to one another.

Neolacon Miwa. The genus was established for three species, formosanus Miwa,
cervinus Erichson, sinensis Candéze ‘and probably others’ in which the lateral
carina of the prothorax almost attains the anterior margin. The length of this
carina, which arises at, or just anterior to the posterior angles of the prothorax,
has been found to vary considerably both intra-specifically and on the two sides of
the same individual. Five years after establishing Neolacon, Miwa (1934 : 14)
placed it in synonymy with Compsolacon (q.v).

Colaulon Arnett. The genus was erected for those Nearctic species ‘which
have hitherto been placed in the genus Lacon [auctt. = Agrypnus as defined in
this work], but which differ from the type-species murinus Linnaeus, in that the
scaly vestiture is sparse, coarse and not coloured and forms regular rows on the
elytra and in which the lateral margin of the prothorax is denticulate’. Since
these characteristics are in my opinion of no value at the generic level (see p. 114),
and as all the Colaulon species known to me possess the diagnostic characteristics of
Agrypnus the genus is here treated as a synonym of Agrypnus.

Cryptolacon Nakane & Kishii. The genus was erected for species differing from

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 117

Compsolacon by the absence of a carina on the posterior angles of the prothorax
and from Colaulon by the absence of tarsal grooves on the propleurae. Kishii
(1961 : 25) reduced Cryptolacon to a subgenus of Colaulon (q.v.).

Sabikikoreus Nakane & Kishi. This subgenus of Agrypnus was erected on
the basis of a number of variable characteristics including the shape of the body,
the presence of tubercles on the pronotum, the propleural tarsal grooves, the lateral]
carinae of the prothorax and the appearance of the terminal abdominal sternite
(see below). Kishii (1964 : 15) raised Sabikikoreus to generic status.

Sagojyyo Kishii. The type-species yuppe Kishii possesses all the diagnostic
characters of Agrypnus. It differs from the type-species, murinus (Linnaeus)
only by such very variable features as the shape of the body, the presence of tarsal
grooves and the presence of nodules on the pronotum.

Ohira, 1968) : 364 and 1969a : 93 uses the erroneous spelling Sagojo.

Archontoides Cobos. The diagnostic characteristic, the sloping humeri, is
associated with the reduction of the wings. The metasternum and scutellum are
also reduced in length. See also Myrmodes, p. 115.

Pyrganus Golbach. The genus was established for eight Central American species
differing from Colaulon in that the lateral margins of the prothorax are not crenate.
The type-species tuspanensis possesses all the generic characteristics of Agrypnus
and differs from the type-species murinus Linnaeus only in the colour and arrange-
ment of the scales clothing the body, a character which is of importance only at
the specific level. It is of interest that in two of the species included in Pyrganus
by Golbach (paleatus Champion and scarrosus Candéze) the lateral margins of the
prothorax are distinctly crenate. L. dwbiuws Candéze, another species included
in the new genus by Golbach, is not congeneric with the remaining species. It
is a true Lacon species.

SEXUAL DIMORPHISM. Ina large number of species it is impossible to distinguish
the sexes by external characteristics. However the present investigation has
shown that the males and females of certain species display differences in the appear-
ance of one or more of the last three visible abdominal sternites and that in other
species the sexes can be separated on the structure of the antennae or tarsi.

Candéze (1874 : 43) was the first to record that in the female of certain species,
the central area of the last visible abdominal sternite is smooth, whereas in the
male the entire surface is punctured. Champion (1894: 263) appears to have
been unaware of Candéze’s discovery because he states that the smooth patch is
typical of the male. Lewis (1896 : 336) accepted Champion’s statement. That
this did not orginate from a printer’s error is shown by the fact that all the specimens
in the BMNH labelled as males by Champion and Lewis are females and vice-
versa. This type of sexual dimorphism is not found in all species, and does not

occur in any Australian species.

_ The appearance of the surface of last visible sternite varies greatly from one
species to another but it is constant within a species. The form of the differentiated
area, where it occurs, ranges from a slightly raised patch with a well defined outline

118 C¢. Ms Fo von" HAVEK

and polished surface without any large punctures through all the intermediate
stages to an ill-defined area in which the punctures are only very slightly less dense
than on the rest of the underside. Species in which the females display this charac-
teristic to a greater or lesser degree occur in the Palaearctic (e.g. murinus (Linnaeus),
cinnamomeus (Candéze)), Oriental (e.g. binodulus (Motschulsky), setiger (Bates),
hispidulus (Candéze)), Ethiopian (e.g. foedus (Candéze)) and Madagascan regions
(e.g. vestitus Klug, cinctupes (Candéze)) and in North America (e.g. vectangularis
(Say)) and Central America (e.g. paleatus (Champion)). The Northern Indian
lutosus (Candéze) is unusual in that both the fourth and fifth visible abdominal
sternites bear a polished patch in the mid line. The Madagascan species sinuatus
(Candéze) differs from all other species known to me in that the last visible
abdominal sternite of the female bears a single enlarged raised puncture from
which a number of short slender scales arise. In appearance this raised puncture
resembles those found on the fourth visible abdominal sternite of certain Australian
species.

Australian Agrypnus species differ from those from other regions in that the
modified areas, where they occur, lie on the fourth, and in one species also on the
third, visible abdominal sternite. A large raised area with a number of punctures
is found in some species such as ursulus (Candéze) and Jaticollis (Candéze). The
polished area is smaller but much more strongly raised and bears only two or three
punctures in stigmosus (Blackburn) and poriginosus (Candéze). A single large
raised puncture, from which several short narrow scales arise, occurs in a number
of species including punctipennis (Candéze) and marmoratus (Candéze). A large
single puncture of this kind is present on both the third and fourth visible abdominal
sternite of adustus (Elston). The function of these pores and patches is unknown.

The antennae of the male are considerably larger and more strongly serrate than
those of the female in the Indian tvuncatus (Herbst) and probably also the
Australian gracilis (Blackburn) of which only the male is known.

Sexual dimorphism of the tarsal segments is known in only one species, obscurus
Fleutiaux from Africa (= abstrusus see p. 120). In this species the fourth tarsal
segment of the male is distinctly lobed while that of the female is only obliquely
truncate.

DISTRIBUTION. Agrypnus species have been recorded from all parts of the
world except South America. The largest number of species has been described
from the Oriental and Australian regions.

BIOLOGY AND HABITS. Hardly anything has been published on the habits of
the adult beetles. Agrypnus murinus (Linnaeus) is generally recorded as living
under stones and in the ground. A number of species have been taken from the
foliage of sandal (Fleutiaux, 1933: 1). A. eucalypti (Blackburn) was found under
the bark of Eucalyptus. Specimens in collections have provided no information
other than having been taken by sweeping, beating or in light traps. Fleutiaux
(1944 : 148) records that Compsolacon species were found together with Octocryptus
coomant near lakes and rivers and in damp places.

A great deal has been written on the Elaterid ‘clicking’ mechanism and the part

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 119

this ability to jump when turned on their backs plays in the life of the beetles.
According to Horion (1949 : 118) murinus, when turned on its back, refuses to
right itself in the accepted Elaterid fashion but defends itself by everting the paired
[? stink] glands lying near the tip of the abdomen. I have found no descriptions
of the behaviour of other Agrypnus species under similar circumstances.

The larvae of a number of species, including murinus have been described and
figured. They live in the ground and in so far as is known they are carnivorous
though some (e.g. murinus) may feed on roots. Van Emden (1945) and Ohira
(1962) contain useful bibliographies of works on Elaterid larvae.

THE SPECIES INCLUDED IN THE GENUS AGRY PNUS

The majority of species now assigned to the genus were described at the time
when mwurinus Linnaeus was believed to be a typical representative, or later the
type-species, of the genus Lacon and are listed under this generic name in the
Schenkling catalogue. After Fleutiaux’s (1926) correction of the catalogue, a
number of species were described as Adelocera. The following 409 species known
to me are now assigned to Agrypnus.

Agrypnus aberdarensis (Fleutiaux) comb. n.

Adelocera (Archontas) aberdarensis Fleutiaux, 1935) : 199.

LECTOTYPE (present designation). KENyA: 9, Kenya, Mais. Forest, Kinangop,
Mts. Aberdare, vers ouest. 2,600m; Mission de l’?Omo, C. Arambourg, P.-A.
Chappius, R. Jeannel 1932—33; aberdarensis Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 22 ex., same locality as lectotype, one with Fleutiaux’s
determination label (MNHN, Paris). 1 9, same locality as lectotype, with
Fleutiaux’s determination label (BMNH). 1 ex., Elgeyo escarpment, 2500 m, with
Fleutiaux’s determination label (MNHN, Paris). 1 ex., Kikuyu escarpment, 2300 m,
without determination label (MNHN, Paris).

Agrypnus abreptus (Candéze) comb. n.

Lacon abreptus Candéze, 1893¢ : 170.
Adelocera abreptus (Candéze) Fleutiaux, 1926 : 96.

The description is based on unrecorded number of specimens from Kanara,
Belgaum and S. Bombay [India] in the Andrewes collection. Selection of a lecto-
type has been postponed until any material extant in the Candéze collection in the
IRSNB, Brussels has been examined.

Syntypes examined. Inp1a: 1 9, co-type, Belgaum P.; Lacon abreptus Cand.
I g, cotype; Belgaum P.; Andrewes Bequest, 1922: 221. 1 9, co-type; Belgaum
P.; 269; Lacon abreptus Cd., Andrewes Bequest, 1922: 221, 1 9; Belgaum N.;
Andrewes Bequest, 1922 : 221. (BMNH).

120 CaM... o Vion JRA EK

Agrypnus abstrusus nom. n.

Tylotarsus (Lobitarsus) [sic] obscurus Fleutiaux, 1935¢ : 94.
Agrypnus obscurus (Fleutiaux) comb. n. [Junior secondary homonym of Agrypnus obscurus
(Fleutiaux, 1934b).]

LECTOTYPE (present designation). 4, Kalimapeza [position unknown] fevrier
(MacArthur); Tylotarsus obscurus Fleut. [Fleut.] (MNHN, Paris).

Paralectotype. KENyA: 4, Brit. E. Africa, S. Masai Res., 10.iv.co13,) [ae
Anderson; Tylotarsus obscurus Fleut. [Fleut.]; Pres. by Imp. Inst. Ent., B.M.
1935 : 95. (BMNH).

Additional material examined. UGANDA: 3 4, 4 9, Gulu, IV—V. 1925 (G. D. H.
Carpenter) (BMNH).

This species is unusual in that the fourth tarsal segment of the male is lobed,
while that of the female is obliquely truncate.

Agrypnus acerbus (Candéze) comb. n.

Lacon acerbus Candéze, 1888 : 670.
Adelocera acerbus (Candéze) Fleutiaux, 1926 : 96.

Holotype. Bhamo, BurMA collected by Fea. MCSN, Genoa.
The generic attribution is based on the description.

Agrypnus acervatus (Candéze) comb. n.
Lacon acervatus Candéze, 1888 : 670.

Adelocera acervatus (Candéze) Fleutiaux, 1926 : 96.

Syntypes: 2 ex., Plapoo [position unknown], Tenasserim april (Fea). ?>MCSN,
Genoa.

The generic attribution is based on the following specimens; BuRMA: 2 9, Carin
Cheba [see p. 275] v.88 (L. Fea), one bearing Candéze’s determination label

[Candéze’s 1891 : 774 locality] (BMNH). 13, 1@Q same locality both with Candéze’s
determination labels (MNHN, Paris).

Agrypnus aculeatus (Candéze) comb. n.

Tilotarsus [sic] aculeatus Candéze, 1857 : 178.
Centrostethus aculeatus (Candéze) Schwarz, 1898c : 414.

LECTOTYPE (present designation). MApAGAscAR: 4, Madagascar; Janson
coll., 1903 : 130; aculeatus [Candéze label stuck to the underside of Janson’s];
Tilotarsus aculeatus Cdze., type ex. coll. de Laferté [Janson] (BMNH).

Agrypnus acuminipennis (Fairmaire) comb. n.

Lacon acuminipennis Fairmaire, 1878 : 109.
Adeloceva acuminipennis (Fairmaire) Fleutiaux, 1926 : 96.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 121

Type-locality. CENTRAL CurNaA, collected by Abbé David. Present location
of material unknown.

The generic attribution is based on the following specimens; 1 2, Museum Paris,
Moupin; Lacon acuminipennis Fairm. [unknown mss.]; costicollis Cand., acumini-
pennis Fairm. [Fleut.]; Type. 1 9, same locality without determination labels
(MNHN, Paris). These specimens may be part of the syntype-series. Thesynonymy
with costicollis Candéze (Fleutiaux, 1962 : 96) has not been confirmed.

Agrypnus acutangulus (Fleutiaux) comb. n.
Compsolacon acutangulus Fleutiaux, 19344 : I5.
LECTOTYPE (present designation). Cuina: 3g, Tchoung-King, Sze Tchouan

[Chung King, Szechwan]; Compsolacon acutangulus Fleut., type [Fleut.} (MNHN,
Paris).

Paralectotype. CHINA: S. Kansu, 28.1x.30 (NR, Stockholm). Not examined.

Agrypnus adelaidae (Blackburn) sp. rev.
Lacon adelaidae Blackburn, 18914 : 505.
LECTOTYPE (present designation): AUSTRALIA: 3, 1948 T [on card mount, see
p- 270]. Australia, Blackburn coll.; Lacon adelaidae Blackb., {Blackb.| (BMNH).

Comparison of the lectotypes has shown that Elston’s (1924: 197) synonymy
with carinulatus Candéze is not justified.

Agrypnus adeloceroides (Candéze) comb. n.

Tilotarsus [sic] adeloceroides Candéze, 1900 : 79 (3).
Lobotarsus adeloceroides (Candéze) Schwarz, 1906 : 30.

The description is based on an unrecorded number of specimens from ‘Congo
francais Benito’ [EQUATORIAL GUINEA].

Type-material: ?>IRSNB, Brussels.

The generic attribution is based on the following specimens, 2 ex., Benito, Congo
Franc., determined by Fleutiaux (MNHN, Paris). These specimens may be part of
the syntype series.

Agrypnus adustus (Elston) comb. n.

Lacon adustus Elston, 1924 : 205.
Lacon adustus Elston, Neboiss, 1956 : 4.

The description is based on an unrecorded number of specimens from AUSTRALIA:
Queensland; Cairns (A. M. Lea), Coen River, Stewart River (W. D. Dodd), Towns-
ville (N. B. Tindale), Malanda (Dr E. Mjéberg).

122 COM, Fo von on AYwnk

Syntype-material (see p. 274): SAM, Adelaide, AM, Sydney, MNV, Melbourne.
Not examined.

The generic attribution is based on a syntype 92: Cairns; Lacon adustus Elston,
co-type (SAM, Adelaide).

Agrypnus aequalis (Candéze) comb. n.

Lacon aequalis Candéze, 1900 : 78.
Adelocera aequalis (Candéze) Fleutiaux, 1926 : 96.

Holotype. Inp1a: g, Kashmir; 902; Lacon aequalis Cand., type [?Cand.];
Andrewes Bequest, 1922-221 (BMNH).

Agrypnus afflictus (Candéze) comb. n.

Lacon afflictus Candéze, 1874 : 68.
Adelocera afflictus (Candéze) Fleutiaux, 1926 : 98.

The description is based on an unrecorded number of specimens from Malacca
and Siam in the Janson and Candéze collections. The Candéze collection in the
IRSNB, Brussels does not contain any specimens from these localities. The only
specimen in that collection with a Candéze determination label is from Borneo.
It is not conspecific with the lectotype designated below. The BMNH collection
also does not contain specimens from the published localities. The only specimen
with Candéze determination label is from Penang. However since this specimen
agrees well with the description and as, during the last century, Malacca was some-
times used as an alternative name for Malaya I believe that the following lectotype
. designation is justified.

LECTOTYPE (present designation). MAtLaAya: 9, Penang [Janson]; Janson coll.
1903.130; afflictus Cdz. [Cand.]; Lacon afflictus Cand. co-type [Gahan] (BMNH).

Agrypnus akidiformis (Candéze) comb. n.
Myrmodes akidiformis Candéze, 1857 : 169.
Holotype. AvusTRALIA: 3, Nov. Holl. Raffles Bay; Janson coll. 1903.130; akidi-

formis Cdze. Type ex. coll. de Laferte [Janson]; Myrmodes akidiformis Cdz.,
Type [Cand.] (BMNH).

Agrypnus alberti nom. n.

Tylotarsus mocquerysi Fleutiaux, 1934b : 59. ;
Agrypnus mocquerysi (Fleutiaux) comb. n. [Junior secondary homonym of Agvypnus mocqueryst
(Fleutiaux, 1932). ]

LECTOTYPE (present designation). MADAGASCAR: 9, Madagascar, Baie
d’Antomgil, A. Mocquerys; mocquerysi Fleut, type [Fleut.] (MNHN, Paris).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 123

Agrypnus albisparsus (Candéze) comb. n.
Tilotarsus [sic] albisparsus Candéze, 1857 : 174.

Syntypes examined. Mapascar: 1 3g, Latimell Ch. Mad. [illegible manuscript];
albisparsus Cand., Type [Fleut.]. The specimen stands over a Chevrolat collection
label; Tilotarsus albisparsus Cand., type Mon. 1, 1857 p. 174, 4. Madagascar
(MNHN, Paris). 1 9, Madagascar, Janson coll. 1903.130; Tilotarsus albisparsus
Cdz. Madag. [Cand.]; Tylotarsus albisparsus Cdze., ex. coll. Candéze [Janson]
(BMNH). 3 3, Madgascar; Janson coll. 1903.130; Tilotarsus albisparsus Cdz.
ex. coll. de Laferté [Janson]. One specimen bears the word ‘Type’ on Janson’s
determination label (BMNH). The absence of Candéze’s determination label is
probably due to Janson (see p. 276).

The designation of a lectotype has been postponed until the specimen in the
MNHU, Berlin, mentioned by Candéze has been examined.

Agrypnus albitactus (Candéz) comb. n.

Lacon albitactus Candéze, 1874 : 55.
Adelocera albitactus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). NIGERIA: g, Old Calabar; Janson coll.
1903.130; albitactus [Cand.]; Lacon albitactus Cand., Old Calabar [Gahan] (BMNH).

Agrypnus albomaculatus (Miwa) comb. n.

Lacon albomaculatus Miwa, 1934 : 68, pl. I, fig. 7.

Type-material. JAPAN: Honshiu, Gifu, vii, 1914 (Y. Miwa). Neither the
number of specimens or their location is recorded. ? Taiwan Agricultural Research
Institute, Taipei.

The generic attribution is based on the description.

Agrypnus albopictus (Candéze) comb. n.

Lacon albopictus Candéze, 1857 : 104.
Adelocera albopictus (Candéze) Fleutiaux, 1926 : 98.

LECTOTYPE (present designation). MaApacascar: 3, Dej. Madagascar [ Janson];
Janson coll. 1903.130; Lacon albopictus Cdze., Cand. [Janson]; Agrypnus albopictus
Dej. cat e coll. Dejean [Janson] (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see p. 276).

Paralectotypes. 1 g, 2 9 with the same labels as the type (BMNH). The
determination label on one female bears the word ‘type’ but no type-designation
has been published. 1 9, 16102 [MNHU, Berlin catalogue number, refers to locality
and collector on following label]; Madagascar Goud.; albopictus Cand., Madag.,
Goudot [Gerstaecker]. 33, Madagascar, Goud. no. 16102 (MNHU, Berlin). The
absence of Candéze’s determination label is probably due to Gerstaecker (see p. 275).

124 CG. M2 Fs von AME

Agrypnus alboscutatus (Candéze) comb. n.

Lacon alboscutatus Candéze, 18936 : 8.

Adelocerva alboscutatus (Candéze) Fleutiaux, 1926 : 96.
The description is based on an unrecorded number of specimens from MADAGASCAR.
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: I ex, St Maria de
Madagascar; alboscutatus Cand., comparé au type [Fleut.] (MNHN, Paris).

Agrypnus alluaudi (Fleutiaux) comb. n.

Centrosthus alluaudi Fleutiaux, 1934b : 61.

LECTOTYPE (present designation). MApAGAscAR: 9, Madagascar, Forét
Tanala, Alluaud, 1901; Museum Paris, Madagascar, Coll. Ch. Alluaud; Centrostethus
alluaudi Fleut., type [Fleut.] (MNHN, Paris).

Agrypnus alternatus (Schwarz) comb. n.

Lacon alternatus Schwarz, 1902a : 311.
Adelocera alternatus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from NEW
GuInEA, Astrolabe Gebirge.

Type material: ?>DEI, Eberswalde.

The generic attribution is based on Schwarz’s comment that the species shows a
similarity to wallacei Candéze.

Agrypnus amplicollis (Boheman) comb. n.

Lacon amplicollis Boheman, 1851 : 414.
Adelocera amplicollis (Boheman) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). SouTH AFRICA: 4, Caffraria; J. Wahlb.;
Type; amplicollis Boh. [Boh.] (NR, Stockholm).

Paralectotypes. 3 9, Caffraria; J. Wahlb. (NR, Stockholm).
The published locality is ‘Habitat in tractibus fluvii Gariepis’. The discrepancy
between the published locality and the label on the specimen is discussed on p. 280.

Candéze (1857: 161) states that Agrypnus granulosus Eschscholtz (1829 : 32)
may be conspecific with amplicollis Boheman. This has not been confirmed.
The description of Agrypnus granulosus is based on an unrecorded number of
specimens discovered by Westermann in ‘Cap. b. sq. [sic]’. The type-material has
not been located. For the history of the Eschscholtz collection see p. 274.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 125

Agrypnus andersoni (Blackburn) comb. n.

Lacon andersoni Blackburn, 1891 : 508.
Lacon andeysoni Blackburn; Neboiss, 1956 : 4.

LECTOTYPE (present designation). AUSTRALIA: g, 644T [on card mount,
see p. 270]; Australia, Blackburn Coll.; Lacon andersoni Blackb. [Blackb.] (BMNH).
The published locality is S. Australia, near Port Lincoln.

Agrypnus angularis (Schwarz) comb. n.

Tilotarsus angularis Schwarz, 1903a : 359

The description is based on an unrecorded number of specimens from Madagascar.
Syntype-material: ?DEI, Eberswalde.

The generic attribution is based on the following specimen; MADAGASCAR; I ex.,
Tilotarsus angularis Schw. Det. par l’auteur. Madag. (MNHN, Paris).

Agrypnus angulicollis (Candéze) comb. n.

Lacon angulicollis Candéze, 18916 : 243.
Adeloceva (Compsolacon) angulicollis (Candéze); Van Zwaluwenburg, 1959 : 352.

Lectotype (designated by Van Zwaluwenburg, 1959 : 352) JAVA: pass (Peontjak)
between Buitenzorg and Preanger, W. Java (Pasteur); Candéze identification
label (RNH, Leiden). Not examined.

Paralectotypes: 2 ex, same locality as the lectotype (RNH, Leiden).

The description is based on three specimens collected by M. J. D. Pasteur in
western Java. Candéze also records that he possessed a large number of specimens
collected in the eastern part of the island. These may be the ones recorded as
being labelled ‘Ex Typis’ by Van Zwaluwenburg.

Agrypnus angusta (Fleutiaux) comb. n.

Adeloceva angusta Fleutiaux, 1942 : I.

LECTOTYPE (present designation). Burma: g, N. E. Burma, Kambaiti,
2000 m., 13/5.1934, R. Malaise; Adelocera angustata Fleut., co-type [Fleut.] (MNHN,
Paris).

The description is based on an unrecorded number of specimens collected by the
Swedish expedition to Burma and British India in 1934.

Agrypnus antiguus (Candéze) comb. n.

Lacon antiguus Candéze, 1857 : 100.
Adelocera antiguus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MADAGASCAR: 2, 16091 [museum catalogue
number, refers to locality and collector on following label]; Madagascar, Goudot;

126 Ce wis Ys won eEvAWGEK

antiguus Cand. Madag. Goud. [Gerstaecker] (MNHU, Berlin). The absence of
Candéze’s determination label is probably due to Gerstaecker (see p. 275).

Paralectotypes: 2 3, Madagascar, Goud. Nr. 16091 (MNHU, Berlin). x 9, Lap.
Madagascar; Janson coll. 1903.130; Lacon antiguus Cdze. Cand., Type e coll de
Laferté [Janson]; globicollis Gory Madag. [? la Ferté Sénectére] (BMNH). The
absence of Candéze’s determination label is probably due to Janson (see p. 276).

Agrypnus apodixus (Candéze) comb. n.

Lacon apodixus Candéze, 1865 : 9.
Adelocera apodixus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). PHILIPPINES: 9, Luzon [Janson]; 394;
Janson coll. ex Candéze, 1903.130; Lacon apodixus Cdze. Type (ex. coll. Cand.) —
[Janson] (BMNH). The absence of Candéze’s determination label is probably due
to Janson (see p. 276).

Agrypnus applanatus (Elston) comb. n.

Lacon applanatus Elston, 1924 : 206.
Lacon applanatus Elston; Neboiss, 1956 : 4.

The description is based on an unrecorded number of specimens from WESTERN
AUSTRALIA.
Syntype-material (see p. 274): AM, Sydney, SAM, Adelaide, NMV, Melbourne.

The generic attribution is based on the following syntypes: 2 9 on one card,
W. Australia, Note Book 318: co-type (SAM, Adelaide).

Agrypnus arbitrarius (Elston) comb. n.

Lacon arbitrvarius Elston, 1924 : 208.
Lacon arbitvavius Elston; Neboiss, 1956 : 5.

The description is based on an unrecorded number of specimens from NORTH-WEST
AUSTRALIA: Derby, Kimberly district (Dr E. M7dberg).
Syntype-material (see p. 274): AM, Sydney, SAM, Adelaide.

Generic attribution is based on a syntype ¢ bearing a co-type label (SAM,
Adelaide).

Agrypnus arctior (Candéze) comb. n.

Lacon arctioy Candéze, 18950 : 56.
Adelocera arctioy (Candéze) Fleutiaux, 1926 : 96.

Holotype. Mapacascar: 9, Madagascar, Diego Suarez 17, Ch. Alluaud 1893;
Museum Paris, Madagascar, Coll. Ch. Alluaud; Type; Lacon arctior Cand. n. sp.
[Cand.]; Cand. det. [Fleut.] (MNHN, Paris).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 127

Agrypnus arenicola Nakane & Kishii
Agrypnus (Sabikikorius) arenicola Nakane & Kishii, 1955 : 3, pl. 2, figs 4, 5.

Holotype. JAPAN: 9, Takarajima, 29.v.1953, S. Ueno leg. Osaka Mus. Nat.
Hist. Not examined. The generic attribution is based on the description.

Agrypnus argentatus (Candéze) comb. n.

Lacon argentatus Candéze, 18936 : 7.
Adeloceva argentatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from MADAGASCAR;
Antananarivo.

Syntype-material: ?>IRSNB, Brussels.

Generic attribution based on Candéze’s comment that the species resembles a
small twrbidus Germar (see p. 224).

Agrypnus argillaceus (Solsky) comb. n.

Lacon argillaceus Solsky, 1871 : 360.
Archontas argillaceus (Solsky) Fleutiaux, 1947 : 280.

The description was based on material collected in Vladivostock in July by
R. Maak and acquired by Solsky. This material has not been located.

The generic attribution is based on material under this name in the BMNH and
MNHN, Paris. This material includes several different species bearing a close
resemblance to one another.

Agrypnus aristatus (Champion) comb. n.

Lacon avistatus Champion, 1894 : 267.
Pyrganus aristatum (Champion) Golbach, 1968 : 198.

Syntype-material: 26 specimens bearing the characteristic part printed, part
Champion manuscript determination labels: B.C.A. Coll. III (x), Lacon aristatus
Ch. Mexico: 1 4, Laf. Vera Cruz, Mex.; Lacon lezeleucei Cand., Cand. e coll.
de Laferté [Janson]. 1 9, GUATEMALA; Coll. Janson. 3 9, Costa Rica; Caché
H. Rogers (BMNH). 1 ex., same locality (MNHN, Paris). 1 g, Costa Rica, Van
Patten. xr 9, Costa Rica; Janson, ex Deyrolle; Lacon lezeleucei [Champ.].
PANAMA: 2 4, 10 Q, V. de Chiriqui, 2-3000ft. Champion (BMNH). 2 ex., same
locality (MNHN, Paris). 2 3, 2 2, Bugata, 800-1500 ft. Champion (BMNH).

Despite the fact that it bears the same labels as the lectotype of lezeleucii Candeze.
I do not believe that the specimen recorded above labelled Jezeleucii Cand. by
Janson is part of the original syntype series of that species. The specimen differs
from the description of Jezeleucii in several ways including the appearance of the
|scales, which are narrow and hair-like, and the posterior angles of the prothorax
which are right-angled and sharp instead of turned outwards and truncate. The

128 COM EF. won ASE K

other specimen labeled lezeleucit was probably misidentified by Champion before
he described aristatus.

The designation of the lectotype is postponed until the material can be studied
in greater detail.

Agrypnus armatus (Candéze) comb. n.
Tilotarsus [sic] armatus Candéze, 1889 : 75 (8).

LECTOTYPE (present designation). MapaGaAscar: 3, Museum Paris, Madagas-
car, Goudot 1834; 4177/34; Tilotarsus armatus Cand., type 89 [Cand.] (MNHN,
Paris).

Agrypnus asper (Candéze) comb. n.

Lacon asper Candéze, 1874 : 55.
Adelocera aspey (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). W. Arrica: 9, Niger source [Janson];
Janson coll. 1903 : 130; Lacon asper Cand., type, Niger source [Gahan]; asper
[Cand.] (BMNH).

Paralectotypes: 2 9, Niger source; Janson coll. 1903 : 130; Lacon asper Cand.
[Gahan] (BMNH).

The published locality is ‘Guinée, Bouches du Niger (Coll. Janson)’. It seems
probable that the discrepancy between the published and label localities arose
when Janson relabelled the specimens.

Agrypnus asperulatus (Candéze) comb. n.

Lacon aspevulatus Candéze, 1878a : 103.
Adeloceva (Compsolacon) asperulata (Candéze); Van Zwaluwenburg, 1959 : 352.

Lectotype (designated by Van Zwaluwenburg, 1959 : 352). NEW GUINEA:
New Guinea, Isola Yule (D’Albertis) (MCSN, Genoa). Not examined.

Paralectotypes. 9 ex., standing beside lectotype (Van Zwaluwenburg, loc. cit.)
(MCSN, Genoa); Not examined. 4 ex., N. Guinea, Isola Yule, T.F.v. 1875; L. M.
D’Albertis; Lacon asperulatus Cand. [unknown mss.] (MNHN, Paris). 4 ex., same
locality, one specimen with Fleutiaux’s determination label and another label
Ann. Mus. civ. Genoa 1878 p. 103. [Fleut.]._ 2 ex., same locality with date iv.1875:
2 ex., same locality with date vi.1875 (MNHN, Paris).

Lacon assus (Candéze) comb. n.

Lacon assus Candéze, 1857 : 145.
Lacon assus Candéze; Neboiss, 1955 : 5.

LECTOTYPE (present designation). AUSTRALIA: sex undetermined, Nov..
Hollandia; Janson coll. ex Candéze, 1903.130; Lacon assus Cdz. N. Holl. [Cand.];
Type ex coll. Candéze [Gahan] (BMNH). The abdomen is missing.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 129

Agrypnus badeni (Candéze) comb. n.

Lacon badeni Candéze, 18786 : LIII(7).
Adelocera badeni (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from
MADAGASCAR.
Syntype-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: 1 ex. Ihosy, Madagasc.;
Adelocera badeni Cand. (Lacon) [Fleut.] (BMNH.).

Agrypnus baibaranus nom. n.

Neolacon formosanus Miwa, 1929 : 235; fig. 2.

Agrypnus formosanus (Miwa) comb. n. [Junior secondary homonym of Agrypnus foymosanus
(Bates, 1886).]

The description is based on an unrecorded number of specimens from Formosa,
Baibara.

Type-material: ?Taiwan Agricultural Research Institute, Taipei.

The generic attribution is based on the description.

Agrypnus bakeri (Fleutiaux) comb. n.
Lacon bakeri Fleutiaux, 1916 : 219.

LECTOTYPE (present designation). PHILIPPINES: 9, Butuan, Mindanao,
Baker; Lacon bakeri Fleut., type, J. Sc. Phil. 1916: 219 [Fleut.] (MNHN, Paris).

Agrypnus basalis (Fleutiaux) comb. n.

Adeloceva (Archontas) basalis Fleutiaux, 19350 : 200.

LECTOTYPE (present designation). KENnyA: 9, Kenya, Mais. forest. Kinangop
Mt. Aberdare vers ouest, 2,600 m.; Museum Paris, Mission de L’Omo, C. Arambourg,
P. A. Chappuis, R. Jeannel, 1932-33; basalis Fleut. type [Fleut.] (MNHN, Paris).

Paralectotypes. 7 ex., same locality as lectotype, two with Fleutiaux’s deter-
mination label (MNHN, Paris). 1 9, same locality was lectotype, determined by
Fleutiaux (BMNH). 1 ex., Plateau de Uasin Gishu; Soy St. II, 1800 m., (MNHN,
Paris).

The metathorax and wings are reduced in length.

Agrypnus beccari (Candéze) comb. n.

Lacon beccavi Candéze, 1880 : 192.
Adeloceva (Compsolacon) beccavi (Candéze) ; Van Zwaluwenburg, 1959 : 352.

Lectotype (designated by Van Zwaluwenburg, 1959 : 325): SUMATRA: Mt. Singalan,
I

130 Cc. M. BF. von HAYEK

Sumatra (MCSN, Genoa). Candéze’s determination label is stuck on a catalogue
card. Not examined.

Paralectotype. (Van Zwaluwenburg, loc. cit.) sex and data not recorded (IRSNB,
Brussels). Not examined.

The generic attribution is based on Candéze’s comment that beccari resembles
Lacon furunculosus Candéze (see p. 157).

Agrypnus benitensis nom. n.

Lobotarsus depressus Schwarz, 1902a : 312.
Agrypnus depressus (Schwarz) comb. n. [Junior secondary homonym of Agrypnus depressus
(Candéze, 1874).]

Syntypes examined. EQUATORIAL GUINEA; I 2 Benito, Congo Franc; Lobotarsus —
depressus Schw. n.sp. [Schw.]: Lobotarsus depressus Schw. type [Fleut.]. 1 9,
same locality without determination label but labelled ‘paratype’. (MNHN, Paris).
1 9, Benito, Congo Franc; Coll. Schwarz; Tylotarsus [unknown mss.] depressus
[Schwarz]; Type (DEI, Eberswalde).

The designation of the lectotype is postponed pending the critical examination
of the material.

Agrypnus bergeali (Girard) comb. n.
Archontas bergeali Girard, 1970 : 29.

Holotype. 3, Mapacascar: Madagascar centre, environs d’Ambohimahasoa,
canton de Tsarafidy, forét d’Ankafina, 1,450m, 12.xi.1963 (P. Viette) (MNHN,
Paris). Not examined.

Paratypes [number not recorded] same locality and date as holotype (NMHN,
Paris and Girard Collection, Laboratoire de Zoologie, Ecole normale supérieure,
Paris). Not examined.

The generic attribution is based on the description.

Agrypnus bidentata (Fleutiaux) comb. n.
Adeloceva (Archontas) bidentata Fleutiaux, 1934) : 53.
The description is based on an unrecorded number of specimens from MApDaA- |
GASCAR: Mont Tsaratanana décembre (Descarpentries) in the author’s collection.
Syntype-material: not located, >MNHN, Paris.

The generic attribution is based on Fleutiaux’s comment that this species resembles
A. turbidus (Germar) and A. antiguus (Candéze).

Agrypnus bidivisus (Candéze) comb. n.

Lacon bidwisus Candéze, 1874 : 58.
Adeloceva bidivisus (Candéze) Fleutiaux, 1926 : 96.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE I31

LECTOTYPE (present designation). NIGERIA: 9, Calabar; bidivisus Murr.
Calab. [Candéze, blue border]; bidivisus Murr. [Cand.]; Lacon bidivisus Cd. det
E. Candéze [IRSNB curatorial label]; Collection E. Candéze (IRSNB, Brussels).

Murray did not describe a species of this name. Candéze based his description
on an unrecorded number of specimens in the Murray collection. No specimens
of this species have been found in the Janson collection in the BMNH (see note on
Murray collection, p. 279). It would appear that in this case Candéze retained the
material submitted to him for determination.

The metasternum is greatly reduced in length and wings appear to be entirely
absent. The fourth tarsal segment is lobed.

Agrypnus bigener (Elston) comb. n.

Lacon bigener Elston, 1924 : 208.
Lacon bigeney Elston; Neboiss, 1956: 5.

The description is based on an unrecorded number of specimens from AUSTRALIA:
Northern Territory, Port Darwin (W. K. Hunt.)

Syntype-material (see p. 274): SAM, Adelaide, AM, Sydney, NMV, Melbourne.

The generic attribution is based on a syntype 9, labelled co-type (SAM, Adelaide).

Agrypnus bimarginatus (Schwarz) comb. n.

Lacon bimarginatus Schwarz, 1908c : 93.
Adeloceva bimarginatus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on a large number of specimens from Madagascar,
Montagnes d’Ambre, received from Rolle, Berlin.

Syntype-material: ?DEI, Eberswalde.

The generic attribution is based on the following specimens: MADAGASCAR: 9,
Mont d’Ambre, determined by Fleutiaux (BMNH). 1 ex., Amber Geb., Coll.
Schwarz, determined by Fleutiaux (MNHN, Paris).

Agrypnus binodulus (Motschulsky)

Lacon binodulus Motschulsky, 1861a : 8.
Agrypnus binodulus (Motschulsky); Kishii, 1964 : 14.

The description is based on an unrecorded number of specimens collected by

: Madame Gaschkevitch in Japan. This material has not been located. It may be
in the Motschulsky collection in the ZMU, Moscow.

The generic attribution is based on the following specimen; JAPAN: g, Ins. Jesso
[Hokkaido]; Janson coll. 1903.130; binodulus Motsch., named by Candéze [Gahan]
(BMNH).

Kishii (1961 : 24; pl. 1, figs 1 and 5) described the subspecies coreanus from
Korea and Tsushima Island. I have not had the opportunity of examining these

132 Ch Me 1B) von HAYEK

specimens but the shape of the aedeagus shown in the figure suggests that these
specimens may not be congeneric with the generally accepted interpretation of
binodulus represented by the material in the BMNH and MNHN, Paris.

The date of the description of binodulus is generally recorded as 1860, the date
on the title-page. Griffin (1936) is of the opinion that the work did not appear
until 1861 and in view of the fact that the introduction on p. 4 bears the date
‘r Nov. 1860’ Griffin’s dating has been accepted.

Agrypnus bipapulatus (Candéze)

Lacon bipapulatus Candéze, 1865 : 11.
Agrypnus bipapulatus (Candéze) Nakane & Kishii, 1955 : 4.
Agrypnus sakishimanus Ohira, 19670 : Ioo.
Agrypnus (Agrypnus) bipapulatus sakishimanus Ohira; Ohira, 1969 : 90.

Lacon bipapulatus Candéze. LECTOTYPE (present designation). 9, CHINA:
China; Janson coll. 1903.130. Lacon bipapulatus Cdz., China [Cand.]; Lacon
bipapulatus Cdze. Type (ex coll. Cand.) [Janson] (BMNH).

Lacon sakishimanus Ohira. Holotype. JAPAN: dg, in Ohira collection.

Paralectotypes. 3 9, Entomological Laboratory, College of Agriculture, Ehine
University and/or Nagao, Sato, Subata and Kishii collections. I have not had the
opportunity of examining this material.

Agrypnus bipunctatus (Schwarz) comb. n.

Lacon bipunctatus Schwarz, 1908a : 82.
A delocera bipunctatus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Madagascar,
Plateau de l’Amdroy [sic], Region de Ambovombe.

Type-material: ? DEI, Eberswalde.
The generic attribution is based on the following specimen; MADAGASCAR: 4,

Plateau de l’Androy, Reg. de Ambovembe; Lacon bipunctatus Schw., comp. au
type [Fleut.] (MNHN, Paris).

Agrypnus bipunctatus (Schwarz) is a junior secondary homonym of Agrypnus
bipunctatus Candéze, 1857 (= Lanelater bipunctatus (Candéze) Arnett, 1952) but
since the two species are not congeneric the junior name does not require replace- |
ment (CZN, Art. 59(b)). |

Agrypnus blackburni (Fleutiaux) comb. n.

Lacon duplex Candéze, 1897 : 9. [Junior primary homonym of Lacon duplex Blackburn, 1892.]
Adelocera blackburn Fleutiaux, 1926: 97. [Replacement name for Lacon duplex Candéze.]

LECTOTYPE (present designation). INDIA: 9, Sikkim, D. Atkins, 88; n.sp.
1893, duplex Cand., Sikkim, m. Genes [Candéze, yellow border]: Collection E. |

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 133

Candéze; Lacon duplex Cand. Det. E. Candéze [IRSNB, curatorial label]. (IRSNB,
Brussels).

The specimen measures 12-4 mm in length compared with the published measure-
ment of II mm.

Agrypnus blairei (Fleutiaux) comb. n.
Adeleocera blaivei Fleutiaux, 1927 : 77.

LECTOTYPE (present designation). NortH VIETNAM: 2, Tonkin, Lac Tho,
Hoa Binh, A. de Cooman; Adelocera blairei Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 2 3, 4 9, same locality as the lectotype, one with Fleutiaux’s
determination label. Laos: 1 9, Pou-Lan, Ht. Mekong 13.5.18; Laos, Luan-Prabang
environs, Vitalis de Salvaza (MNHN, Paris).

Agrypnus brachychaetus (Kollar) comb. n.

Lacon brachychaetus Kollar, 1844 : 506.
Adeloceva brachychaetus (Kollar) Fleutiaux, 1926: 9.

The description is based on an unrecorded number of specimens from KASHMIR,
presumably collected by Huegel.
Type-material: >NM, Vienna. (Horn, 1935 : 110).

The generic attribution is based on the following specimens, which may be syntypes
2 §, Kashmir; Museum Paris, Collection Leon Fairmaire 1905; Lacon brachychaetus
[?Fairm.] Fairm. det. [Fleut.] (MNHN, Paris).

Agrypnus brevis (Candéze) comb. n.

Lacon brevis Candéze, 1857 : 156.
Colaulon breve [sic] (Candéze) Golbach, 1968 : 1098.

LECTOTYPE (present designation). Mrxico: 3, Yucatan; Coll. Janson, ex
Candéze, Lacon brevis Cdz. Yucatan 9 [Cand.]; Lacon brevis Cand. ex coll. Candéze
[Gahan] (BMNH).

Paralectotypes. 1 9, brevis Cand. Yucatan [Champ]; Coll. Janson, ex Candéze
(BMNH).

I ex., Yucatan; L. brevis [Cand.]; Collection Chevrolat; Lacon brevis Cand,
typ. Mon. [Fleut.] (MNHN. Paris).

Agrypnus brightensis (Blackburn) comb. n.

:
: Lacon brightensis Blackburn, 18914 : 503.
| Lacon brightensis Blackburn, Neboiss, 1956 : 5.
LECTOTYPE (present designation). AUSTRALIA: g, 651 AL. T. [on card mount,
see p. 270]; Australia, Blackburn Coll.; Lacon brightensis Blackb. [Blackb.]. (BMNH)

134 iC. Mis Py vow EA Ek

Paralectotypes: An unrecorded number of specimens in the AM, Sydney and
SAM, Adelaide (Neboiss, 1956 : 5).

The published locality is Victoria; near Bright, under bark of Eucalyptus.

The wings and metasternum are much reduced in length.

Agrypnus brunneipennis (Candéze) comb. n.

Lacon brunneipennis Candéze, 1857 : 133.
Adelocera brunneipennis (Candéze) Fleutiaux, 1926 : 96.

Holotype. N. Inp1A: g, N. India [Janson]; Janson coll. ex Candéze 1903.130;
Lacon brunneipennis Cdz., N. Indoun [sic, Cand.]; Lacon brunneipennis Cand. ex
coll. Candéze [Gahan] (BMNH).

The published locality is ‘Indes-Orientales’, see p. 271.

Agrypnus caffer (Candéze) comb. n.

Lacon caffey Candéze, 1881 : 7.
Adelocera caffey (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). S. Arrica: Caffrarie; n.sp. caffer Cdz.,
Caffr. [Candéze, yellow border]; Collection E. Candéze; ex type; Lacon caffer Cand.,
det. E. Candéze [IRSNB curatorial label} (IRSNB, Brussels).

Agrypnus calamitosus (Candéze) comb. n.

Lacon calamitosus Candéze, 1874 : 100.
Pyrganus calamitosus (Candéze) Golbach, 1968 : 108.

The description is based on an unrecorded number of specimens from Mexico:
Oajaca [sic] in the Sallé collection.

The generic attribution is based on the following specimen: MExico: 2, Playa-
Vincente; Mexico, Salle Coll; 3, B.C.A. Col. (1) Lacon calamitosus Cand. [Champ];
Lacon calamitosus Cand. Sallé coll. 1420 [unknown mss. the number refers to Sallé
mss. Catalogue] (BMNH).

Playa-Vincente is 110 km N.E. of Oaxaca. Champion (1895 : 266) regarded this
specimen as the type, though he was aware of the discrepancy in the localities.
The specimen is not recorded as a type in the Sallé mss. catalogue in the BMNH.
Lectotype designation has been postponed until a search can be made for Candéze’s
material in the IRSNB, Brussels.

Agrypnus caliginosus (Boisduval) comb. n.

Adelocerva grisea Guérin, 1830, pl. 2, fig. 7. [Nomen oblitum.]

Adelocera caliginosa Boisduval, 1835 : 98.

Elater caliginosus Boisduval, 1835 : 105. [Synonymized with Lacon caliginosus (Guérin) by
Candéze, 1857 : 144.]

Adelocera caliginosa Guérin, 1838 : 68.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 135

Lacon caliginosus (Boisduval) Germar, 1840 : 261.
Lacon caliginosus (Guérin); Neboiss, 1956 : 5.

This is the species which all workers, including Neboiss (1956 : 5), author of the
most recent catalogue of Australian Elateridae, have credited to Guérin, generally
with the date 1832.

The date 1832 for the ‘Voyage autour du Monde...sur la Corvette... La
Coquille pendent les années 1822, 1823, 1824 et 1825’, volume 2, part 2, in which a
description of Adelocera caliginosa Guérin appears on page 68, was given by Hagen
(1863 : 310) and accepted by many subsequent bibliographers. However, according
to Kirkaldy (1902 : 317) Hagen did not see this work, which though dated 1830
on the title page, bears the date 1838 at the end of the Avant-propos (p. xii) and
contains references in the text to works published after 1832 (Bequaert, 1926 : 186).
There can be no doubt that volume 2, part 2 of the text was published in 1838.
The plates illustrating the insects described in the text were published between
May 1830 and December 1831 (Guérin, 1838 : 271, Sherborne and Woodward,
1906 : 338). Plate 2 was published on 25th November, 1830.

The ready acceptance of Hagen’s dating of Guérin’s work was probably due
to the fact that Boisduval (1835) gives references not only to the plates illustrating
the voyage of the Coquille (published 1830-31, see above) but also to the text.
For example on page 115 under Lyctus parallelus which is not illustrated by Guérin,
Boisduval refers to Guerin, Voy. de la Coquille, Zool. p. 72, the page on which the
description appeared in 1838. According to Sherborn and Woodward (1906 : 336)
this ‘is accounted for by the new widely recognized fact that the contemporary
writers, working on the different French Voyages, then in course of compilation,
often exchanged the proofs of their several works’. Boisduval, probably included
Guérin’s species because it was his intention (1835 : iii) to furnish his reader with
‘un espéce de faune entomologique de l’Oceanie’.

Adelocera caliginosa Boisduval. The description is presumably based on material
collected at ‘Hobart Town et a la terre de van Diemen [TASMANIA] in the course of
a voyage around the world by the Astrolabe under the command of J. Dumont
D’Urville. This collection should be in the MNHN, Paris (Boisduval, 1832 : 11,
Horn & Kahle, 1935 : 63) but the most careful searches have failed to discover
the material.

The fact that Boisduval refers to Guérin, Voy. de la Coquille, pl. ii, fig. 7 suggests
that he believed that his Tasmanian caliginosus was identical with the Australian
species depicted by Guérin, 1830 (with the name grisea, subsequently changed to
caliginosa, see below). Since the species at present known as Lacon caliginosus
(Guérin) is one of the few species which is fairly common in S.E. Australia and
Tasmania, I suggest that a neotype of Adelocera caliginosa Boisduval be selected
when suitable Tasmanian material belonging to the species at present known as
Lacon caliginosus (Guérin) is found in the MNHN, Paris.

Elater caliginosus Boisduval. LECTOTYPE (present designation). AUSTRALIA:
2, Dej. Nov. Holl. [Janson]; Agrypnus caliginosus Dej. Cat. ex. coll. Dejean [ Janson];
Lacon caliginosus Cand. [Janson] Guer. [Gahan]; Janson coll. 1903 : 1330 (BMNH).

136 C. Mi. Fy von HAW EK

The absence of a Dejean determination label (Horn & Kahle 1935: pl. 24, no 45)
is probably due to Janson, who may also have removed Boisduval’s label, if it ever
existed.

Why Boisduval failed to recognize that his Agrypnus caliginosa and Elater
caliginosus are identical remains a mystery. It is possible that Guérin’s comment
(1838, Avant-propos) that Boisduval worked fast and carelessly is fully justified.

Adelocera caliginosa Guérin. The description is based on material from Port
Jackson [Sydney, N.S.W.] collected in the course of a voyage around the world
by the Coquille, commanded by Captain L. I. Duperry. This collection should be
in the MNHN, Paris (Boisduval, 1832 : ii, Horn a Kahle 1936 : 258) but, once again,
the most careful searches have failed to discover the material. Examination of the
material standing as Lacon caliginosus (Guérin) in the BMNH and MNHN, Paris
has shown that there is a general agreement concerning the identity of this species —
I suggest that a neotype of this species be selected when suitable material becomes
available in the MNHN, Paris.

The figure (pl. 2, fig. 7) illustrating Guérin’s description of Adelocera caliginosa
was published (see p. 135) on 25th November, 1830. The legend of the plate gives
the name as Adelocera grisea. The reason why Guérin changed the name of his
species, adopting that used by Boisduval, may be found in the Avant-propos
(1838 : xi and 271). There he points out that Boisduval was not justified in dis-
regarding the generally accepted practice of adopting a name published in conjunc-
tion with a figure or description. However, in order to show the lengths to which
he was prepared to go to avoid accusations of confusing the synonymy, he proposed
to accept Boisduval’s new names for his figures. It is this magnanimous action
together with the erroneous date on the title page that misled many workers into
believing that Guérin’s text was published before that of Boisduval.

Adelocera grisea Guérin, 1830. This is an available name in that it fulfills the
provision of the International Code of Zoological Nomenclature, 1961. As the
material on which the figure is based cannot be found (see above) and as I.C.Z.N.
Article 75(6) forbids the designation of a neotype for a name not in general use,
I am of the opinion that, in the interest of stability an application should be made
to have Adelocera grisea Guérin, 1830 placed on the Official Index of Rejected
Names. :

Candéze (1857: 161) remarks that Agrypnus cribrosus Eschscholtz (1829),
described from Hawaii, bears some resemblance to caliginosus Guérin. The
type-material of crvibrosus has not been located but the description, size and
distribution suggests that cribrosus may be conspecific with Lacon modestus
(Boisduval). See also p. 72.

GERMAR’S INTERPRETATION OF Lacon caliginosus BOISDUVAL

Germar ascribed caliginosus to Boisduval, Faune de l’Oceanie, Elater caliginosus.
It may be assumed that he is referring to the species described by Boisduval, 1835 :
105. Germar records the locality of E. caliginosus as Van Diemansland [Tasmania] |

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 137

in place of Boisduval’s Nouvelle Holland [Australia]. This may be due to Germar
confusing the localities of Adelocera caliginosa Boisduval and Elater caliginosus
Boisduval or because his interpretation of the species was based on Tasmanian
material identified by means of Boisduval’s description.

Neither the BMNH, DEI, Eberswalde nor NMHU, Berlin, contain specimens
from Tasmania determined by Germar. The BMNH contains a male and female
from the Schaum collection. The male bears a determination label believed to
be in Germar’s handwriting (Horn & Kahle, 1935; pl. 37, no. 36); caliginosus De}.
Boisd., N. Holl. The female bears the number 1022 (referring to the manuscript
catalogue of the Schaum collection) and a small blue triangular label with the
letters ‘Vd’. These letters may indicate Van Diemansland. The specimens
agree well with Germar’s description though the male is smaller (11-5 mm) and
the female larger (13-6 mm) than Germar’s published measurement of 6 lines
(=12:-8mm). Both specimens are conspecific with the lectotype of F. caliginosus
Boisduval.

Despite the fact that the specimen with Germar’s determination label originates
from Australia, I am of the opinion that it represents Germar’s interpretation of
E. caliginosus Boisduval.

Agrypnus candezei (Fleutiaux) comb. n.

Lacon candezei Fleutiaux, 1895a : 685.
Adelocera candezei (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). NortH VIETNAM: 9, Tonkin, Florentin;
Lacon candezei type [Fleut]; Lacon inops Cand., Cand. det. [Cand.]; L. candezei
Fleut. Ann. Soc. ent. Fr. 1894 : 685 [Fleut]. (MNHN, Paris).

Paralectotypes. 3 ex., same locality as the lectotype, but without Fleutiaux’s
determination labels (MNHN, Paris).
The published locality is Lang-Son.

Agrypnus canescens (Candéze) comb. n.

Lacon canescens Candéze, 1897 : 9.
Archontas canescens (Candéze) Cobos, 1970 : 133.

LECTOTYPE (present designation). W. Arrica: 9, n. sp. 1893 [sic] canescens
Cand., Congo [Candéze, blue border]; Lacon canescens det. E. Candéze [IRSNB
curatorial label]; Collection E. Candéze; Ex. Typis (IRSNB, Brussels).

The published locality is Ogoowé [Gabon].

Agrypnus cariei (Fleutiaux) comb. n.

Lacon caviei Fleutiaux, 1920b : 490.
Adelocera cariei (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Mauritius: 3, Ie Maurice, Butte 4
l'Herbe, Carie, Juillet 1900; Lacon cariei Fleut. type [Fleut.] (MNHN, Paris).

138 CAM I) vont HAY Ec

Paralectotypes: 1 ex., Mt. C. de Garde, U. d’Emmerez; Museum Paris, Madagascar
[crossed out] Coll. Ch. Alluaud 1904; Lacon cariei Fleut. [Fleut.]. 1 g, Ile Maurice,
Curipepe Levieux 1895; Lacon desjardinsi Cand.; Museum Paris, Coll. P. Carié
1914; Lacon cariei Fleut. [Fleut.]. 1 g Ile Maurice, Curipepe, Carié 1897; Museum
Paris, Coll. P. Carié 1914 (MNHN, Paris).

Agrypnus carinicollis (Schwarz) comb. n.

Lacon carinicollis Schwarz, 1908a : 83.
Adelocera carinicollis (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Madagascar.
Type-material: ?>DEI, Eberswalde.

The generic attribution is based on the following specimen; MADAGASCAR:
Foréts Nord, Ft. Dauphine, Alluaud; carinicollis Schw. [Fleut.] (MNHN, Paris).

Agrypnus carinulatus (Candéze) comb. n.

Lacon carinulatus Candéze, 1857 : 152.
Lacon carinulatus Candéze; Neboiss, 1956 : 6.

LECTOTYPE (present designation). AUSTRALIA: 9, Deyr. N. Holl.; Janson
coll. 1903.130; carinulatus Cdz., type [Cand.]; Lacon carinulatus Cand. type (e
coll. Deyrolle) [Janson] (BMNH).

Paralectotype: sex undetermined, Deyr. N. Holl.; Janson coll. 1903.130; Lacon
carinulatus Cdze. Cand. e. coll. Deyrolle [Janson] (BMNH). The abdomen is
missing.

The metasternum is reduced in length and the wings are short.

Agrypnus castaneipennis (Candéze) comb. n.

Lacon castaneipennis Candéze, 1956 : 55.
Adelocera castaneipennis (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected in
Tananarive, MADAGASCAR by Alluaud.
Type-material: ?>IRSNB, Brussels and/or Alluaud collection, MNHN, Paris.

The generic attribution is based on a specimen from Madagascar, Mt. Poraka,
determined by Fleutiaux (MNHN, Paris).

Agrypnus cervinus Erichson comb. n.

Elater (Agrypnus) cervinus Erichson, 1834 : 230.
Adelocera cervinus (Erichson) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). PHILIPPINES: g, I61II; cervinus Er.*
Manilla, Mayen [Gerstaecker] (MNHU, Berlin).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 139

Agrypnus cinctipes (Germar) comb. n.
Tylotarsus cinctipes Germar, 1840 : 247.

LECTOTYPE (present designation). MApAscAR: g, Janson coll. ex Schaum,
1903.130; cinctipes Dj. [plus illegible word, crossed out] Klug, Madagascar [believed
to be Germar’s handwriting] (BMNH).

Germar records that he received this species determined as Agrypnus cinctipes
Dej. from Dupont. Although there is no incontrovertible proof that this is Germar’s
original specimen, I believe it has a better claim than the two males in the MNHU,
Berlin bearing the number 16083. One also bears a Gerstaecker label: cinctipes
Germ., cuspidatus Cand., Madag. Goudot. Both are conspecific with the lectotype.

Agrypnus cinerascens (Candéze) comb. n.

Lacon cinerascens Candéze, 1879a : 103.
Adelocera cinerascens (Candéze) Flautiaux, 1926 : 96.

Holotype. NeEw Guinea: Yule Is., collected by D’Albertis, >MCSN, Genoa.

The generic attribution is based on the following specimen: 1 ex., N. Guinea Mer.
Hula, Gemaio 1891, L. Loria; cinerascens Cand. [Fleut.]; Cand., Ann. Mus civ.
Genoa 1892, p. 801 [Fleut.] (Candéze, 1891, locality) (MNHN, Paris).

Agrypnus cineraceus (Elston) comb. n.

Lacon cineraceus Elston, 1927 : 359.
Lacon cineraceus Elston; Neboiss, 1956 : 6.

The description is based on an unrecorded number of specimens from AUSTRALIA:
Queensland; Bowen (A. Simson), Normanton (R. Kemp).

Syntype material: (see p. 274) SAM, Adelaide, AM, Sydney.

The generic attribution is based on the following syntype: 9, Normanton (R.
Kemp) (SAM, Adelaide).

Agrypnus cinnamomeus (Candéze) comb. n.

Lacon cinnamomeus Candéze, 1874 : 76.
Lacon cinnamomeus var. a, Candéze, 1874 : 76.
Adelocera cinnamomeus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). USSR: 3, Siberia or.; Janson coll. 1903.

130; Lacon cinnamomeus Cand., type. E. Siberia [Gahan]; Lacon n.sp. (cinnamomeus)
| [Janson]; Lacon argillaceus Solsky [Gahan] (BMNH). The absence of Candéze’s

| determination label is probably due to Janson (see p. 276).

_ Paralectotypes. 2 3g, Siberia or.; Janson coll. 1903.130, one specimen bears
an additional label; Lacon cinnamomeus Cand. E. Siberia [Gahan] (BMNH).

A. cinmnamomeus var. a. LECTOTYPE (present designation). Korea: 3,

140 CoM. Fo Von HAWEK

Corea, Vladimir Bay A. Adams; Janson coll. 1903 : 130; cinnamomeus var. [Cand.]
(BMNH).

The published locality is ‘Vladimir Bay dans la Corée. Coll. Janson.’ As far 4
as I can discover Vladimir Bay (43°52’N., 135°30’E.) is in Russia, about 200 miles
north of the Korean border.

Fleutiaux’s (1918a : 191) synonymy with argillaceus Solsky, 1870 has not been
confirmed.

Agrypnus coarctatus (Candéze) comb. n.

Lacon coarctatus Candéze, 1874 : 70.
Adelocera coarctatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from HIMALAYA
in the Janson collection.
Type-material: not found in BMNH; ?lost.

The generic attribution is based on the following specimen; 3, BuRMaAgH, deter-
mined by Schwarz (BMNH).

Agrypnus coctus (Candéze) comb. n.

Lacon coctus Candéze, 1874 : 66.
Adelocera coctus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Burma: 3, Rangoon; Janson coll. 1903.
130; L. coctus Cdz. type [Cand.]; Lacon coctus Cand. type [Gahan] (BMNH).

Paralectotypes. 1 3, Rangoon; Janson coll. 1903.130. L. coctus Cdz. type,
[Cand.]. 1 g, Rangoon; Janson coll. 1903.130; L. coctus Cdz. [Cand.]. 3 g, 1,
Rangoon, Janson coll., 1903.130 (BMNH).

Agrypnus coenosus (Hope)

Elatey coenosus Hope, 1831 : 25.

Agrypnus coenosus (Hope) Hope, 1843a : 63.

Lacon nepalensis Candéze, 1874 : 72. Syn. n.
Adelocera nepalensis (Candéze) Fleutiaux, 1926 : 96.

Elater coenosus Hope. LECTOTYPE (present designation). NEPAL: 3, Nepal;
coensus Hope 4034; Elater coenosus Hope, type Grays Z. Misc. [Gahan];
Hardwicke Bequest (BMNH).

Lacon nepalensis Candéze. LECTOTYPE (present designation). NEPAL: 9, |
Nepal; Janson coll. 1903.130; nepalensis Cdz. [Cand.]; Lacon nepalensis Cand., |
type [Gahan] (BMNH).

Paralectotype. 2, Hardwicke, Nepal; Janson coll. 1903.130 (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 141

Agrypnus cognatus (Van Zwaluwenburg) comb. n.

Compsolacon cognatus Van Zwaluwenburg, 1957 : 8, fig. 2a, 36.
Adelocera (Compsolacon) cognatus Van Zwaluwenburg; Van Zwaluwenburg, 1966 : 298.

Holotype. E. CARroLines: 9, No. 62925. (USNM, Washington). Allotype:
2, same locality (BPBM, Honolulu). Not examined.

The generic attribution is based on the description and figures.

Van Zwaluwenburg (1966: 298) believed this species to be a secondary junior
homonym of A delocera cognatus Candéze, 1892, and proposed a new name A. nesiotes.
As the species are not congeneric (see also Lacon cognata (Candéze) p. 61), Van
Zwaluwenburg’s new name is unnecessary.

Agrypnus collisus (Candéze) comb. n.

Lacon collisus Candéze, 1891a : 772.
Adelocera collisus Candéze; Fleutiaux, 1926 : 96.

The description is based on two syntypes from BurMA: Montagnes de Carin,
[Karin] detroit des Cheba (900-1100) metres, decembre, Fea collection.

Syntypes: ?>MCSN, Genoa.

The generic attribution is based on Candéze’s comment that the species resembles
colonicus Candéze and taciturnus Candéze. Both these species are now attributed
to Agrypnus.

Agrypnus colonicus (Candéze) comb. n.

Lacon colonicus Candéze, 1881 : 8.
Adelocera colonica (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Cochin-
China [S. Vietnam] in the Mniszech collection.

Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen; Laos: 1 g, Ht. Laos;
Lacon colonicus Cand. [Fleut.] (BMNH).

Miwa (1934: 186) places colonicus in synonymy with Paralacon taciturnus
(Candéze) but Fleutiaux (1947 : 263) treats colonicus and taciturnus as separate
species of Adelocera.

In my opinion the specimen on which the generic attribution of colonicus is
based is conspecific with the syntypes of taciturnus but examination of the type-
material of colonicus is necessary before the synonymy can be confirmed.

Fleutiaux (1947 : 264) suggests that Paralacon kawamurae (Miwa, 1929 : 230)
may be asynonym of this species. This cannot be confirmed until the type-material
of kawamurae has been located and examined.

142 CM ES vio) LASEK

Agrypnus colorata (Fleutiaux) comb. n.
Adeloceva (Archontas) colovata Fleutiaux, 19346 : 53.

LECTOTYPE (present designation). _MApDAGASCAR: 3g, Madagascar, Baie
d’Antongil, A. Mocquerys; colorata Fleut., type [Fleut.]. (MNHN, Paris).

Agrypnus commutabilis (Elston) comb. n.

Lacon commutabilis Elston, 1924 : 207.
Lacon commutabilis Elston, Neboiss, 1956 : 6.

The description is based on an unrecorded number of specimens from AUSTRALIA:
Northern Territory: Port Darwin (W. K. Hunt), coll. of Dr E. W. Furguson; North
West Australia: Derby (W. D. Dodd), Noonkanbah (Dr E. Mjoberg).

Syntype-material (see p. 274): AM, Sydney, SAM, Adelaide, NMV, Melbourne.

The generic attribution is based on the following specimen; AUSTRALIA: 9, Port
Darwin, labelled cotype (SAM, Adelaide).

Agrypnus comptus (Candéze) comb. n.

Lacon comptus Candéze, 1874 : 65.
Adelocera comptus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected by
Muhot and de Castelnau in ‘Siam; Camboge’ in the Janson, Saunders and Candéze
collections.

Syntypes examined. CAMBODIA: I g, Camboja; Shantbon [position unknown];
Janson coll. 1903.130; comptus Cdz. [Cand.]; Lacon comptus Cand. [Gahan].
THAILAND: I 9, Pachebon [?Phet Buri, see Adelocera mouhoti p. 35], Muhot; Janson
coll. 1903.130, ex. coll. Saunders; comptus Cdz. [Cand.] (BMNH).

Designation of a lectotype is postponed until any specimens extant in the Candéze
collection in the IRSNB, Brussels have been examined.

Agrypnus consobrinus (Candéze) comb. n.

Lacon consobrinus Candéze, 1857 : 125.
Adeloceva consobrinus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Inp1A: 9, Dej. India [Janson]; 285; Janson
coll. 1903.130; Lacon consobrinus Cdze, Type e coll. Deyrolle [Janson]; consobrinus
[Cand] (BMNH).

The published locality is ‘Indes-Orientales’, see p. 271.

Agrypnus consors (Candéze) comb. n.

Lacon consors Candéze, 1874 : 64.
Adelocera consors (Candéze) Fleutiaux, 1926 : 96.

¥

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 143

LECTOTYPE (present designation). INDIA: sex not determined, Madras;
Janson coll. 1903.130; consors Cdz. [Cand.]; Lacon consors Cand. type [Gahan].
(BMNH). The abdomen is missing.

Agrypnus conspiciendus (Elston) comb. n.

Lacon conspiciendus Elston, 1924 : 206.
Lacon conspiciendus Elston; Neboiss, 1956 : 6.

The description is based on an unrecorded number of specimens from AUSTRALIA:
Northern Territory, King River.

Type-material (see p. 274): SAM, Adelaide, AM, Sydney.

The generic attribution is based on the following specimen: AUSTRALIA: 9, Finke
R. Gaus. (Dr H. Basedon); Lacon conspiciendus Elst. Finke R., C.A. (SAM, Adelaide).
The specimen bears a close resemblance to Agrypnus akidiformis (Candéze).

Agrypnus conspurcatus (Candéze) comb. n.

Lacon conspurcatus Candéze, 1895) : 56.
Adeloceva conspurcatus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MApAGASCcAR: 3g, Madagascar, Diego
Suarez, Ch. Alluaud, 1893; Museum Paris, Madagascar, Coll. Ch. Alluaud; Type:
Lacon conspurcatus Cand. n. sp. [Cand.]; Cand. det. 1895 [Fleut.] (MNHN, Paris).

Paralectotypes: II ex., same locality aslectotype. One bears a label: conspurcatus

Cand., ex type. Ann. Belge 1895: 56. (MNHN, Paris). 1 9, same locality as
lectotype, with Fleutiaux’s determination label (BMNH).

Agrypnus cordicollis (Candéze)

Lacon cordicollis Candéze, 1865 : 9.
Agrypnus cordicollis (Candéze) Ohira, 1957 : 7.

The description appears to be based on a single specimen from Japan in the
ZMU, Helsinki. Not examined.
The generic attribution is based on the following specimen: 1 J, JAPAN, determined
by Janson (BMNH).

Agrypnus cordipennis (Candéze) comb. n.

| Lacon cordipennis Candéze, 1874 : 99.
| Lacon cordipennis Candéze; Neboiss, 1956 : 6.

LECTOTYPE (present designation). AUSTRALIA: 9, Port Albany, N. N. Holl;
Janson coll. 1903.130; cordipennis Cdz. type [Cand.]; Lacon cordipennis Cand.
type, Albany [Gahan] (BMNH). The metasternum and wings are reduced in
} den

144 Cu Ma ES vonvHAvEK

Agrypnus costicollis (Candéze) comb. n.

Lacon costicollis Candéze, 1857 : 116.
Ayrchontas costicollis (Candéze) Fleutiaux, 1947 : 282.

Holotype. Candéze based his description on a single specimen from ‘Indes
Orientales’ (see p. 171) submitted by Deyrolle. The specimen is not in the BMNH.
?IRSNB, Brussels.

The generic attribution is based on the following specimen: INpDIA: Q, India;
Lacon costicollis Cand. [Janson]; (BMNH).

Agrypnus costipennis (Germar) comb. n.

Lacon costipennis Germar, 1848 : 181.
Lacon yilgarnensis Blackburn, 1892b : 289. Syn. n.
Lacon costipennis Germar; Neboiss, 1956 : 6.

Lacon costipennis Germar. LECTOTYPE (present designation). AUSTRALIA:
3d Adelaide, Germ. Nr. 16137 [museum catalogue number = Adelaide, Germar]
(MNHU, Berlin).

Paralectotypes: 1 g, 16137; Adelaide Germ.; 1 g, Adelaide Germ. Nr. 16137
(MNHU, Berlin).

The specimens stand beside a light blue label with Gerstaecker handwriting:
costipennis Germ.* Cand. Adelaide Germ. The absence of Germar’s determination
label is probably due to Gerstaecker (see p. 275).

Lacon yilgarnensis Blackburn. LECTOTYPE (present designation). AUSTRALIA:
9, 4294T. Yilgarn [on card mount, see p. 270]; Lacon yilgarnensis Blackb. [Blackb.]
(BMNH).

The description is based on an unrecorded number of specimens from Yilgarn,
a goldfield in W. Australia, of which Southern Cross is the centre.

Agrypnus crenatus (Klug) comb. n.

Elatery (Agrypnus) crenatus Klug, 1833 : 66 (1834 : 154).
Lacon terrenus Germar, 1840 : 262. [Synonymized by Candéze, 1857 : 103.]
Adeloceva crenatus (Klug) Fleutiaux, 1926 : 96.

Elater (Agrypnus) crenatus Klug. LECTOTYPE (present designation). MaApA-
GASCAR: 4, I6100 [museum catalogue number = Madagascar, Goudot]; Madagascar
Goudot (MNHU, Berlin).

Paralectotype: g, 16100; Madagascar Goudot (MNHU, Berlin). The specimens
stand beside a blue Gerstaecker label: crenatus Cand.,* Elat. crenatus Klug, L.
terrenus Germ. Madag. Goud.

Lacon terrenus Germar. LECTOTYPE (present designation). MADAGASCAR:
g, 1019; Janson coll. ex Schaum, 1903.130; terrenus Germ., Madag. sp. propr.
[Schaum] (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 145

Germar based his description of tevrenus on an unrecorded number of Madagascan
specimens received from Dupont. As Candéze (1857 : 104) records that he received
‘le L. terranus de Germar’ from Schaum, it is reasonable to assume that Schaum
relabelled the specimen while it was in his collection. There are no specimens
bearing Germar’s determination labels in the ZMHU, Berlin or DEI, Eberswalde.

For the history of the Germar collection see p. 275.

Germar (1840 : 263, 440) appears to have based his interpretation of crenatus
Klug on Madagascan material collected and determined by Dupont. I have not
been able to trace any of Dupont’s (see p. 274) specimens bearing Germar’s determi-
nation label. Germar’s own collection was broken up (see p. 275). Neither the
MNHU, Berlin nor the DEI, Eberswalde collections possess specimens with Germar’s
determination labels. The BMNH manuscript catalogue of the Schaum collection
lists three specimens of crenatus Klug, and the collection contains the three specimens
recorded below. These specimens agree well with Germar’s description, in which
he makes especial mention of the crenate lateral margin of the elytra, a distinctive
feature not shown by crenatus Klug. Despite the absence of Germar’s determina-
tion label, I have no doubt that the following three specimens which appear to
belong to an undescribed species, formed part of, if not all, of the material examined
by Germar.

MADAGASCAR: I 4, Janson coll. ex Schaum, 1903.130; Dp. [manuscript on small
blue triangular label, may indicate that this is Dupont’s Madagascan material]
(BMNH). 1 dg, Janson coll. ex Schaum, 1903.130 (BMNH). 1 9, Janson coll. ex.
Schaum, 1903.130; crenatus Klg. [Cand.] (BMNH).

Agrypnus crenicollis (Ménétriés) comb. rev.

Elater crenicollis Ménétriés, 1832 : 156.

Agrypnus crenicollis (Ménétriés) Faldermann, 1835 : 166.

Lacon crenicollis (Ménétriés) Germar, 1840 : 265, pl. 1, fig. I.

Adelocera (Compsolacon) crenicollis (Ménétriés) Van Zwaluwenburg, 1966 : 289.

The description is based on a large number [‘Je l’ai trouvé en grand quantité . . .]
of specimens from: Montagnes de Schiste, devant l’Elbouz [U.S.S.R.}. Fleutiaux
(1947 : 47) states that the type [sic] is in the ZI, Leningrad.

The generic attribution is based on specimens from the Caucasus standing under
this name in the BMNH and MNHN, Paris. There is a general agreement concern-
ing the identity of this species.

GERMAR’S INTERPRETATION OF Lacon crenicollis MENETRIES. Germar gives
only a short description of crenicollis from S. Russia without recording whether
he possessed any specimens. I have not been able to trace any specimens deter-
mined by Germar. As Germar’s figure agrees well with the specimens determined
jas crenicollis known to me, I have assumed that Germar identified the species
correctly.

K

146 CM. Bs von, HAVE TK

Agrypnus curtus (LeConte) sp. rev., comb. n.

Adelocera curtus LeConte, 1853 : 491.
Lacon curtus (LeConte) Candéze, 1857 : 156.
Adeloceva curtus LeConte; Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). U.S.A.: 9, brick red disk [= Southern
States, see p. 277]; curtus 2 (MCZ, Harvard). Length: 10-7 mm.

Paralectotype. 9, brick red disk; Type, 2379 [MCZ curatorial label]; Lacon
curtus Lec. [LeC.] (MCZ, Harvard). Length: 9-6 mm.

The published locality is Georgia. As LeConte records the length as -43 [inches
= II-3 mm] the larger specimen, which is also more the complete, of the two speci-
mens in the LeConte collection has been designated as the lectotype. Why the
paralectotype bears a label with the name Lacon in LeConte’s handwriting is un-
known. LeConte may have replaced his original label with one to correspond with ~
Candeéze’s Monograph.

Arnett (1952: 118) and also Golbach (1968: 198) place curtus in synonymy
with rectangularis Say, 1825. Since the type of vectangularis is lost (see p. 204)
and the synonymy cannot be confirmed, curtus is here treated as a good species.

Arnett (loc. cit.) comments that the species displays a range of variation in the
puncturation and the size of the interstriae of the elytra. This occurs in the material
standing as cuvtus in the BMNH collection. These specimens also display a range
of variation in the length of the wings. Some resemble the type-material in that
the wing attains the base of the fifth abdominal sternite, while in others the wings
barely attain the posterior margin of the third abdominal sternite.

LeConte states that ‘this is the Adelocera curtus of the Dejean catalogue’. Four
specimens (I g, 3 9, Am. Bor.) from the Dejean collection in the BMNH agree well
with the lectotype, except that they are slightly smaller in size (g—10-5 mm.) and
have shorter wings.

Agrypnus cuspidatus (Klug) comb. n.

Elatey (Conoderus) cuspidatus Klug, 1833 : 66 (1834 : 154).
Tilotarsus [sic] boildieur Candéze, 1857 : 176. [Synonymized by Candéze, 1874 : 108.]
Centrostethus cuspidatus (Klug) Schwarz, 1898c : 414.

Elater (Conoderus) cuspidatus Klug. LECTOTYPE (present designation).
MADAGASCAR: 4, 16086 [museum catalogue number = Madagascar, Goudot];
Madagasc. Goudot (MNHU, Berlin). The specimen stands beside a _ blue
Gerstaecker label: cuspidatus Klug, boildieui Cand. Madag. Goud. The absence of
Klug’s label is probably due to Gerstaecker (see p. 275).

Tilotarsus boildieut Candéze. Holotype ¢, Tilotarsus boildieui Cand., type |
[Gahan]; Coquerel; Tilotarsus cuspidatus Klug (Rev.) boildieui Cdze (Mon.) ex |
coll. de Laferté [Janson]; Janson coll. 1903.130 (BMNH). The absence of a Candéze’s
determination label is probably due to Janson (see p. 276).

The published locality is Madagascar.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 147

Agrypnus cylindripennis (Fleutiaux) comb. n.

Adelocera (Archontas) cylindripennis Fleutiaux, 1934b : 57.

LECTOTYPE (present designation). Mapacascar: 9, Ambositra; Adelocera
(Archontas) cylindripennis Fleut. type [Fleut]. (MNHN, Paris).

Agrypnus davidi (Fairmaire) comb. n.

Lacon davidi Fairmaire, 1878 : 109.
Lacon argillaceus var. davidi Fairmaire; Fleutiaux, 1918 : 191.
Archontas argillaceus var. davidi (Fairmaire) Fleutiaux, 1947 : 279.

The description is based on an unrecorded number of specimens from CENTRAL
CHINA, collected by the Abbé David. The present location of the material is
unknown.

The generic attribution is based on the following specimen which may be a syn-
type; 9, China, A. David; L. davidis [sic] Frm. [?Fairm.]: Museum Paris, Coll.
R. Oberthur, 1952 (MNHN, Paris).

Examination of material in the BMNH and MNHN, Paris has shown that a
number of species of very similar appearance are included under argillaceus. In
my opinion argillaceus Solsky, cinnamoneus Candéze and davidi Fairmaire should
be treated as separate species until such time as the type-material of all three
species can be examined.

Agrypnus dealbatus (Candéze) comb. n.

Lacon dealbatus Candéze, 1882 : 13.
Lacon dealbatus Candéze; Neboiss, 1956 : 7.

The description is based on an unrecorded number of specimens from C. York
(Australia, Queensland].

Type-material: IRSNB, Brussels. Neboiss’s statement that the type is in the
IRSNB, Brussels cannot be accepted as a lectotype designation. Not examined.

The generic attribution is based on the following specimen which is probably
a syntype; AUSTRALIA: 9, C. York; dealbatus Cdz. [Cand.] (MNHN, Paris).

Agrypnus deboulayi (Candéze) comb. n.

: Lacon deboulayi Candéze, 1874 : 89.
: Lacon duboulayi Candéze; Neboiss, 1956: 7. [Unjustified emendation.]

LECTOTYPE (present designation). AUSTRALIA: g, Nikol Bay/N.W. N. Holl;
Janson coll. 1903.130; L. deboulayi Cdz. type [Cand.]; Lacon deboulayi Cand.
type, Nikol Bay [Gahan]. Lacon n.sp. debilicornis [Janson] (BMNH).

The metasternum is very short and the wings reduced to short flaps.

148 C. M.. Fo von HAVEK

Agrypnus decoratus (Candéze) comb. n.

Lacon decovatus Candéze, 1882 : 6.
Adeloceva decovatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Madagascar.
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: MADAGASCAR: Q,
Nossi Be, Coll. L. Fairmaire; decoratus Cand. [Cand.] (BMNH).

Agrypnus defectus (Candéze) comb. n.

Lacon defectus Candéze, 1888 : 669.
Adelocera defectus (Candéze) Fleutiaux, 1926 : 96.

The description is based on two specimens from BuRMA: Tenasserim; Thagata,
avril, Kawkareet, mai, collected by Fea, 1885-7.

Syntypes: ?>MCSN, Genoa.

The generic attribution is based on the description.

Agrypnus delesserti (Candéze) comb. n.
Lacon delesserti Candéze, 1882 : 9.
Adelocera delesserti (Candéze) Fleutiaux, 1926 : 96.
The description is based on an unrecorded number of specimens from INDIA:
Neelgherries, collected by Delessert.
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen; INp1a: H. L. Andrewes
Nilgiri Hills; Lacon delesserti Cdze. (BMNH).

Agrypnus denticollis (Fleutiaux) comb. n.

Lacon denticollis Fleutiaux, 1918) : 237.
Adelocera denticollis (Fleutiaux) Fleutiaux, 1926 : 98.

LECTOTYPE (present designation). KENyA: 3, Museum Paris, Afrique Orient. |
Angl. Sud. du lac Rudolphe, Entre le Chemin de fer et le lac. Maurice Rothschild |
1906; Lacon denticollis Fleut. type 1918 [Fleut]; = tellini Fleut. [Fleut] (MNHN,
Paris).

Paralectotype. I ex, same locality as lectotype, but without determination
label (MNHN, Paris).

The synonymy with fellini Fleutiaux, 1903 is unpublished and has not been
confirmed.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 149

Agrypnus depressus (Candéze) comb. n.

Lacon depressus Candéze, 1874 : 77.
Adelocera depressus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). KoreA (NorTH): 9, Corea, Chosan, A.
Adams; Lacon coll. 1903.130; Lacon depressus Cdz. [Cand.]; Lacon depressus
Cand., type Chusan [Gahan] (BMNH).

Agrypnus desjardinsii (Candéze) comb. n.

Lacon desjardinsii Candéze, 1857 : 143.
Adelocera desjardinsii (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). g, Mauritius; Janson coll. ex Candéze,
1903.130; Lacon desjardinsii Cdz. 31. Maur. [Cand.]; Lacon desjardinsii Cand. ex.
coll. Candeze [Gahan] (BMNH).

Paralectotypes. MAuriITIUS: 34, Dej., Ile de France [Janson]; Janson coll.
1903.130; Lacon desjardinsii Dej. Cdze. Cand. [Janson]; Agrypnus desjardinsii
Dej. Cat. e coll. Dejean [Janson]. 1 g, Dej: Ile de France [Janson]; Janson coll.
1903.130; Lacon desjardinsii Dej. Cand. type [Janson]; desjardinsii Dej. Cat. e
coll. Dejean [ Janson} (BMNH).

Candéze states that he received specimens from de la Ferté Sénectére, Mniszech
and Boheman and also refers to Agrypnus desjardinsii Dej. Cat. ed. 3, p. 99. I
have not been able to trace any specimens determined by Candéze from these
collections (see notes on collections consulted, p. 270).

Agrypnus desquamatus (Candéze) comb. n.

: Lacon desquamatus Candéze, 1857 : 124.
Adelocera desquamatus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation): 2 Deyr. India [Janson]; 284; Janson coll.
1903.130; Lacon desquamatus Cdze. Type e coll. Deyrolle [Janson] (BMNH).
The absence of Candéze’s determination label is probably due to Janson (see p. 276).

The published locality is ‘Indes-Orientales’, see p. 271.

Agrypnus deyrollei nom. n.

Lacon coenosus Candéze, 1857 : 114.

Adelocera coenosus (Candéze) Fleutiaux, 1926 : 96.

Agrypnus coenosus (Candéze). comb. n. [Junior secondary homonym of Agrypnus coenosus
(Hope, 1831).]

LECTOTYPE (present designation): 9, Deyr. India [Janson]; 228; Janson coll.
1903.130; L. coenosus [Cand.]; Lacon coenosus Cdze. Type e coll. Deyrolle [Janson]
| (BMNB#).

The published locality is ‘Indes Orientales’, see p. 271.

150 C. M. F. von HAYEK

Agrypnus dilaticollis (Fleutiaux) comb. n.
Adelocera (Archontas) dilaticollis Fleutiaux, 1934b : 50.
LECTOTYPE (present designation). 9, MaApaGAscAR: Madagascar, entre

Soanierana et Foulpointe, Museum Paris; Adelocera (Lacon) [Fleut]; dilaticollis
Fleut, type [Fleut.] (MNHN, Paris).

Agrypnus discedens (Candéze) comb. n.

Lacon discedens Candéze, 1878a : tot.
Adelocera discedens (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Ternate,
[INDONESIA] (Beccari).

Type-material: ?>MCSN, Genoa and/or IRSNB, Brussels.

The generic attribution is based on the following specimen: I 3, 5 ex., TERNATE,
L. Laglaize. One bears a label; Lacon discedens Cand. type [Cand.] (MNHN,
Paris).

It is possible that these specimens are part of the syntype series, though how
they come to be in the MNHN, Paris, isa mystery. I have not been able to discover
whether Beccari visited Ternate. However Léon Laglaise, a French traveller and
collector who was in touch with the Italian collectors in the region at the time,
is known to have visited the island in 1877 (Salvadori, 1878 : 310).

Agrypnus divaricatus (Candéze) comb. n.

Lacon divaricatus Candéze, 1865 : 12.
Lacon divaricatus Candéze; Neboiss, 1956 : 7.

LECTOTYPE (present designation). AuSTRALIA: 2, N. Holl. Victoria, C.
Cdze. [Janson]; Lacon divaricatus Cdz. Victoria [Cand.]; Lacon divaricatus Type
(ex coll. Cand.) [Janson] (BMNH).

Candéze records the locality, under victoriae, as ‘dans les environs de Melbourne’.

Agrypnus dorcinus (Candéze) comb. n.

Lacon dorcinus Candéze, 1875 : CXIX.
Adelocera dorcinus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from the |
Philippines; Bojol [sic. ? Bohol].

Type-material: ?I[RSNB, Brussels.

The generic attribution is based on the following specimens; PHILIPPINES: 6 ex.,
Dapitan, Mindanao, Baker; dorcinus Cand. [Fleut.] (Fleutiaux, 1916 : 219, locality) |
(MNHN, Paris).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 151

Agrypnus duplex (Blackburn) comb. n.

Lacon duplex Blackburn, 1891a : 506.
Adelocera (Compsolacon) duplex (Blackburn) Van Zwaluwenburg, 1959 : 353.

LECTOTYPE (designated by Van Zwaluwenburg, loc. cit.). AUSTRALIA: T. 1347V
[on card mount, see p. 270]; Australia, Blackburn coll.; Lacon duplex Blackb.
[Blackb.] (BMNH).

Paralectotypes (recorded by Neboiss, 1966: 6) AM, Sydney, SAM, Adelaide.
The published locality is Victoria.

Agrypnus elliensis (Candéze) comb. n.

Lacon elliensis Candéze, 1865 : Io.
Adelocera elliensis (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). CEYLON: J, Ceylon; Janson coll. 1903.130
Lacon elliensis Cdz. Ceylon [Cand.]; Lacon elliensis Cdze. Type ex coll. Cand.
[Janson] (BMNH).

Agrypnus ellipticus (Candéze) comb. n.

Lacon ellipticus Candéze, 1857 : 135.
Lacon propinquus Candéze, 1857 : 135. [Synonymized by Candéze, 1891 : 22.]
Adelocera ellipticus (Candéze) Fleutiaux, 1926 : 96.

Lacon ellipticus Candéze. The description is based on an unrecorded number
of specimens from ‘Himalaya’ submitted by Schaum. The BMNH does not possess
any specimens determined by Candéze, nor is the species recorded in the manuscript
catalogue of the Schaum collection. I have been unable to locate the type-material.
It has probably been lost (see history of the Schaum collection p. 280).

The generic attribution is based on the following specimen: 1 3, Muree [INDIA.
N. Punjab] determined by H. E. Andrewes (BMNH).

Lacon propinquus Candéze. The description is based on an unrecorded number
of specimens from ‘Indes Orientales’ (see p. 271) in the Schaum collection. I have
been unable to trace this material (see history of the Schaum collection, p. 280).
The synonymy is provisionally confirmed on the basis of a male from Rangoon,
determined by Janson (BMNH).

Agrypnus elstoni (Neboiss) comb. n.

Lacon conspiciendus Elston, 1927 : 353. [Junior secondary homonym of L. conspiciendus
Elston, 1924 : 206.]
Lacon elstoni Neboiss, 1956: 7. [Replacement name for L. conspiciendus Elston, 1927.]
The description is based on an unrecorded number of specimens from QUEENS-
LAND: Brisbane and NorTH-wEsT AUSTRALIA: Kimberley District (Dr E. Mjoberg).

Type-material (see p. 274): SAM, Adelaide, AM, Sydney.

152 CaM. BY Von EUAMETe

The generic attribution is based on Elston’s comment that the species could be
confused with marmoratus (see p. 181).

Agrypnus eucalypti (Blackburn) comb. n.

Lacon eucalypti Blackburn, 1891a : 507.
Lacon eucalypti Blackburn; Neboiss, 1956 : 7.

LECTOTYPE (present designation). AUSTRALIA: g, Type; 321, [see p. 250]
Lacon eucalypti Blackb. [Blackburn] (BMNH).

The published type-locality is S. Australia, near Port Lincoln, under bark of
Eucalyptus.

Agrypnus eximus (Candéze) comb. n.

Lacon eximus Candéze, 1857 : 101.
Adeloceva eximus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MApDaAGAscAR: 9, Laf. Madagascar; Madag.
Goudot; Janson coll. 1903.130; Lacon eximus Cdze. Cand. type e coll. de Laferté
[Janson] (BMNH). The absence of Candéze’s determination label is probably
due to Janson (see p. 276).

Paralectotypes: 1 9, Madascar; Janson coll. ex Candéze, 1903.130; Lacon eximus
Cdz. Mad. Mnz. [Cand.] (BMNH). 1 9, Madagascar; eximus Cdz. Mad. coll. Mn.

[Cand., yellow border]; Lacon eximus Cand. det E. Candéze [IRSNB, curatorial
label] (IRSNB, Brussels).

Agrypnus fairmairei (Candéze) comb. n.

Lacon fairmairei Candéze, 1889 : 6.
Adelocera faiymairei (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unknown number of specimens from Madagascar
submitted by Fairmaire.
Type-material: ?IRSNB, Brussels and/or MNHN, Paris. See p. 271.

The generic attribution is based on the following specimen; MADAGASCAR: I Q,
Madag. determined from description by Gahan (BMNH).

Agrypnus fallax (Fairmaire) comb. n.

Lacon fallax Fairmaire, 1903 : 202.
Adelocera fallax (Fairmaire) Fleutiaux, 1926 : 98.

LECTOTYPE (present designation). MaApaAcGascar: 2, Madag. Perrier [Fairm.]
Type; Museum Paris, Madagascar, Coll. Perrier de la Bathie 1906; Lacon fallax
Madag. [Fairm.] (MNHN, Paris).

Paralectotypes: 5 ex., Museum Paris, Madagascar, coll. Perrier de la Bathie,

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 153

1906. One specimen is labelled ‘fallax, ex type’. the others lack determination labels.
I ex., Museum Paris, Madagascar, Coll. Leon Fairmaire 1906. Lacks determination
label (MNHN, Paris).

Agrypnus falsarius (Candéze) comb. n.

Lacon falsarius Candéze, 1874 : 60.
Adelocera falsarius (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). CEYLON: 9, Ceylon; Janson coll. 1903.130;
L. falsarius Cdz. [Cand.]; Lacon falsarius Cand. type, Ceylon [Gahan]; L. caelibatus
n.sp. Ressemble a Velliensis mais bien distincte, Fossettes tarsale trés distinctes
sur le pro. et mesothorax, elytres avec des stries de gros pointes L. caelibatus mihi
{Cand.] (BMNH).

Candéze did not publish a description of caelibatus.

Agrypnus farinensis (Blackburn) comb. n.

Lacon farinensis Blackburn, 1900 : 49.
Adeloceva (Compsolacon) farinensis (Blackburn): Van Zwaluwenburg 1959 : 293.

Lectotype (designated by Van Zwaluwenburg, 1959 : 293, see p. 9). AUSTRALIA:
6, 6710 Lyndh. T. [see p. 270] Australia, Blackburn Coll; Lacon farinensis Blackburn
[Blackb.] (BMNH).

Paralectotypes (recorded by Neboiss, 1961 : 6). SAM, Adelaide.

Agrypnus farinosus (Candéze) comb. n.

Tilotarsus [sic] farinosus Candéze, 18954 : 48.
Lobotarsus farvinosus (Candéze) Schwarz, 1898a : 131.

The description is based on 3 specimens from ToGo submitted by Kraatz.
Syntypes: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: g, Toco, Conradt,
determined by Fleutiaux (MNHN, Paris).

Agrypnus fastigiatus (Schwarz) comb. n.

Lacon fastigiatus Schwarz, 1898b : 185.
Adelocera fastigiatus (Schwarz) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). TANZANIA: 9, Usamb. Weise; Coll. Schwarz;
Type; fastigiatus Schw. [Schwarz] (DEI, Eberswalde).

Paralectotypes. 2 9 with locality, collection and type labels as lectotype, 1 9,
Usambara; Coll. Schwarz; Type. (DEI, Eberswalde).

154 C. M. F. von HAYEK

Agrypnus fatuus (Candéze) comb. n.

Lacon fatuus Candéze, 1874 : of.
Lacon fatuus Candéze; Neboiss, 1956 : 8.

Holotype. AUSTRALIA: g, 38153 [Fry catalogue number = data on next two
labels]; De Boulay; N. W. Holl. N. West; Fry coll. 1905.100; Lacon fatuus Cdz.
type [Cand.]; Lacon fatuus Cand. type, Austr. N. W. [Fry] (BMNH).

Agrypnus feneriva (Fleutiaux) comb. n.

Adelocera (Archontas) feneriva Fleutiaux, 19340 : 53.

LECTOTYPE (present designation). MADAGASCAR: 9, Fénérive; feneriva Fleut.,
type [Fleut.] (MNHN, Paris).

Agrypnus feralis (Candéze) comb. n.

Lacon fervalis Candéze, 18916 : 243.
Adelocera feralis (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from W. SUMATRA,
Mandeling [position unknown, ?Mandelang R., 1°04’S., 102°42’E.], in the RNH,
Leiden. Not examined.

The generic attribution is based on the description.

Agrypnus fergusoni (Elston) comb. n.

Lacon fergusoni Elston, 1927 : 350.
Lacon fergusoni Elston; Neboiss, 1956 : 8.

The description is based on an unrecorded number of specimens from Queensland:
Bowen (A. Simpson) and Townsville (E. W. Ferguson).

Syntype-material (see p. 274), AM, Sydney, SAM, Adelaide.

The generic attribution is based on the following specimen: AUSTRALIA: 2, Bowen,
Queensland, A. Simpson; cotype (SAM, Adelaide).

Agrypnus ferrugineus (Candéze) comb. n.

Lacon ferrugineus Candéze, 1874 : 87.
Lacon ferrugineus Candéze; Neboiss, 1956 : 8.

LECTOTYPE (present designation). AUSTRALIA: g, 36820 [Fry catalogue
number = data on next label] Nov. Holl. N. West; Fry Coll. 1905.100; Lacon
ferrugineus Cand., Austra N. West, Type [Fry]; L. ferrugineus [Cand.] (BMNH).

Paralectotype. 9, W. AusTRALIA, Nikol Bay; Janson coll. 1903 : 130; L.
ferrugineus Cdz. [Cand.]; Lacon ferrugineus Cand., co-type [Gahan] (BMNH).

The published localities are: Australie nord-occidentale: Nickol-Bay (Coll. Fry
et Jans.).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 155

Both specimens differ from the description in that a slight depression for the
accommodation of the posterior tarsi is present on the metasternum.

Agrypnus ferruginipes (Fleutiaux) comb. n.
Adelocera (Archontas) ferruginipes Fleutiaux, 1932a@ : 50.

Holotype. Mapacascar: 3, Diego Suarez; Adelocera (Archontas) ferruginipes
Fleut., Type [Fleut.] (MNHN, Paris).

Agrypnus fex (Candéze) comb. n.

Lacon fex Candéze, 1874 : 64.
Adelocera fex (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Malacca
[MALaya] in the Janson collection. This material should be in the BMNH, but I
have been unable to find it.

The generic attribution is based on the following specimen: 9, no locality, L. fex
Cand., Cand. det. [? Fleut.) (MNHN, Paris).

Agrypnus fibrinus (Candéze) comb. n.

Lacon fibrinus Candéze, 1865 : 9.
Adelocera fibyinus (Candézez) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from SUMATRA
in the RNH, Leiden. Not examined.

The generic attribution is based on the following specimen: @ [tentative identifi-
cation, contents of abdomen lost], Sumatra [Janson]; Muller; Janson coll. 1903.130;
Lacon fibrinus Cdz. Sumatra, Sn [Cand.]; Lacon fibrinus Cdze. Type, ex Coll.
Cand. [Janson] (BMNH). This may be one of the original syntype series which
was retained by Candéze (see p. 271) and acquired by Janson.

Agrypnus fictus (Candéze) comb. n.

Lacon fictus Candéze, 1868b : LIII (7).
Adelocera fictus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Madagascar.
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: 9, MADAGASCAR:
Coll. L. Fairmaire; fictus Cand. [Cand.]; Comp. type Fleut.] (MNHN, Paris).

Agrypnus flavescens (Candéze) comb. n.

Lacon flavescens Candéze, 1874 : 66.
Adelocera flavescens (Candéze) Fleutiaux, 1926 : 96.

156 C. M. F. von HAYEK

LECTOTYPE (present designation). CEYLON: g, Ceylon; Janson coll. 1903.130;
flavescens Cdze., Type [Cand.]; Lacon flavescens Cand., Type [Gahan]; Lacon
sericans ? [Janson] (BMNH).

Agrypnus foedus (Candéze) comb. n.

Lacon foedus Candéze, 1857 : 123.
Lacon alboseminatus Kolbe, 1889 : 126. Syn. n.
Adeloceva foedus (Candéze) Fleutiaux, 1926 : 96.

Lacon foedus Candéze. LECTOTYPE (present designation). SENEGAL: 4,
Senegal; Janson coll. ex Candéze, 1903.130; Lacon foedus Cdz. Seneg. {Cand.];
Lacon foedus Cand., ex coll. Cand. [Gahan] (BMNH).

Lacon alboseminatus Kolbe. LECTOTYPE (present designation). ZAIRE: dg,
Kwako bis Kimpoko [4°12’S., 15°33’E. Eastern shore of Stanley Pool]. Kongo,
R. Buttner; Lacon alboseminatus [Kolbe] (MNHU, Berlin).

Agrypnus formosanus (Bates)

Lacon formosanus Bates, 1866 : 348.
Agrypnus formosanus (Bates) Ohira, 1966 : 216.

LECTOTYPE (present designation). Formosa: 9, Ins. Formosa; Janson coll.
1903.130; Lacon formosanus Bates, Formosa [Bates]; Lacon formosanus Bates,
Type (ex coll. Bates) [Janson] (BMNH).

Paralectotype. dg, Ins. Formosa; Janson coll. 1903.130; Lacon formosanus
Bates (ex coll. Bates) [Janson] (BMNH).

Agrypnus foveicollis (Cobos) comb. n.
Lobotarsus foveicollis Cobos, 1970 : 137.
Holotype. CoNGo (BRAZZAVILLE): 6, Sibiti Congo. xii. 1964; Museum Paris,

Mission A. Descarpentries et A. Villiers; Prolobotarsus foveicollis nov. sp. Holotypus
[Cobos] (MNHN, Paris).

Up to the present the generic name Prolobotarsus has not been published.

Agrypnus frenchi nom. n.

Lacon squalescens Blackburn, 1892b : 289.
Agrypnus squalescens (Blackburn) comb. n. [Secondary junior homonym of Agrypnus
squalescens (Fairmaire, 1871).]

The description is based on an unrecorded number of specimens from N. Queens-
land in the French collection (see p. 275). There is no justification for Neboiss’
(1961 : 6) statement that the type is in the BMNH.

The generic attribution is based on the following syntype. AUSTRALIA: 4, 6673

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 157

[see p. 270] N. Qu. T.; Lacon squalescens Blackb. [Blackb.] (BMNH). The species
is not conspecific with Agrypnus marmoratus (Candéze).

Agrypnus fuliginosus (Candéze) comb. rev.

Lacon fuliginosus Candéze, 1865 : Io.
Agrypnus (Sabikikoreus) fuliginosus (Candéze) Nakane & Kishii, 1955 : 3, pl. 2, fig. 5.
Adelocera (Sabikikoreus) fuliginosus (Candéze) Ohira, 1959 : 65.

LECTOTYPE (present designation). JAPAN: 9, Japan, C. Cdz. [Janson]; Janson
coll. ex. Candéze 1903.130; Lacon fuliginosus Cdz. Japan [Cand.]; Lacon fuliginosus
Cdze., Type (ex coll. Cand.) [Janson] (BMNH).

Agrypnus fulvastra (Fleutiaux) comb. n.
Adelocera (Lacon des Auteurs) fulvastva Fleutiaux, 19404 : 91.

LECTOTYPE (present designation). TrBet: 2, Vrionatong, Thibet, Le Moult>
Paris XIII; Adelocera (Lacon) fulvastra n.sp. Fl., Type [Fleut.] (MNHN, Paris).

Agrypnus fulvisparsus (Candéze) comb. n.

Tilotarsus fulvisparsus Candéze, 1874 : 109.

LECTOTYPE (present designation). GABON: 9, Gabon; Janson coll. 1903.130;
T. fulvisparsus Cdz., Type [Cand.]; Tylotarsus fulvisparsus Cdze. type ex coll.
Doné [Janson] (BMNH). Candéze’s (1891 : 27) synonymy with Agrypnus nubilus
(Candéze) has not been confirmed. (see p. 191).

Agrypnus furunculosus (Candéze) comb. n.

Lacon furunculosus Candéze, 1857 : 134.
Adelocera furunculosus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from ‘Des Indes
orientales’ (see p. 271) received from Schaum. I have not been able to locate these
specimens (see histories of the Candéze and Schaum collections p. 271 and p. 280).

The generic attribution is based on the following specimen: 9, furunculosus Cdz.
[Cand.]; Lacon furunculosus Cand., named by Candéze [Gahan]; scutellatus ??
flancs du prothorax sans sillons obliques [unknown mss.] (BMNH). The specimen
lacks a locality label.

Agrypnus fuscus (Fabricius) comb. n.

_ Elatey fuscus Fabricius, 1801 : 228.

Adeloceva fuscus (Fabricius); Van Zwaluwenburg, 1959 : 351.

Lectotype (designated by Van Zwaluwenburg, 1959 : 351). g (not ?9 as stated
by Van Zwaluwenburg), fuscus [Fabricius] (UZM, Copenhagen, Kiel Collection).

158 CoM. Eo von cHAMEBKK

The description is based on material collected in AMBoINA by la Billardiére.

Latreille (1829 : 451) included fuscus Fabricius in Adelocera. In the same year
Eschscholtz (1829 : 32) transfered it to Melanotus and every subsequent worker
has followed suit, listing fuscus Fabricius under Melanotus or one of its synonyms.
The confusion seems to have arisen when Candéze (1860 : 336) synonymized E.fuscus
Fabricius with Cvatonychus fuscus Erichson (1841: 106). The description of
C. fuscus Erichson leaves no doubt that it is not conspecific with E. fuscus Fabricius.
Erichson’s species is probably the Melanotus fuscus (Fabricius) of authors and in
collections.

Agrypnus gabonensis nom. n.

Tilotarsus [sic] veductus Candéze, 1882 : 14.

Lobotarsus veductus (Candéze) Schwarz, 1906 : 30.

Agrypnus veductus (Candéze) comb. n. [Junior secondary homonym of Agrypnus veductus
(Candéze, 1878).]

The description is based on an unrecorded number of specimens from GABON.
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on a specimen from Gabon standing over this
name in the MNHN, Paris.

Agrypnus gazagnairei (Candéze) comb. n.

Tilotarsus [sicl gazagnaivei Candéze, 1889 : 74 (9).
Centrostethus gazagnaivei (Candéze) Schwarz, 1898c : 414.

LECTOTYPE (present designation). MApaAGAscAR: 3g, Madagascar, Museum

Paris, Goudot, 1834; 4177; Lacon detalus Cat.-Mus.; Tilotarsus gazagnairei Cand.
[?Cand.] Candéze vid. [unknown mss.| (MNHN, Paris).

Agrypnus geminatus (Candéze) comb. n.

Lacon geminatus Candéze, 1857 : 149, pl. 2, fig. 5.
Lacon geminatus (Candéze) Neboiss, 1956 : 8.

The description was based on an unrecorded number of specimens from Nouvelle-
Hollande [Australia]. Since Candéze does not record the location of the material
it is almost certain that it is in the BMNH (see history of the Candéze collection,
p. 271). I do not know on what evidence Neboiss (loc. cit.) bases his statement
that the type is in the IRSNB, Brussels. Pending further investigation the BMNH
specimen is here treated as a syntype.

Syntype examined. AUSTRALIA: 9, N. W. Holl., C. Cdze.; Janson coll. ex Candéze;
Lacon geminatus C. N. Austr. Th. [Cand.]; Lacon geminatus Cdze., ex coll. Cand.
[Janson] ? Type [Gahan] (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 159

Agrypnus gibberosus (Candéze) comb. n.

Lacon gibberosus Candéze, 1889 : 71 (5).
Adelocera gibberosus (Candéze) Fleutiaux, 1926 : 96.

The description was based on an unrecorded number of specimens from Mada-
gascar in the MNHN, Paris. I have been unable to find this material. The
specimen recorded below, as Fleutiaux’ labels imply, may be part of Candéze’s
original series.

The generic attribution is based on the following specimen standing as gibberosus
Candéze: 3, MADAGASCAR, Goudot 1834; ? Type ?; type ? ayant perdu son etiquette
[Fleut.] (MNHN, Paris).

Agrypnus gibbus (Candéze) comb. n.

Lacon gibbus Candeze, 1882 : Io.
Lacon gibbus Candéze; Neboiss, 1956 : 8.

The description is based on an unknown number of specimens from Cleveland
Bay, Australia, from the de Monchicourt [Horn & Kahle, 1936: 181, via Candéze
to IRSNB, Brussels], and Mniszech (see p. 278) collections. According to Neboiss
(loc.cit.) the type is in the IRSNB, Brussels, but since Candéze clearly has more
than one specimen and did not designate a type, a formal lectotype designation is
required. I have not seen the type-material in the IRSNB, Brussels.

The generic attribution is based on the following specimen: AUSTRALIA: I 9, L.
gibbus Cand. [Cand.]; Aust. Sept., ex Oberthur (MNHN, Paris).

Agrypnus glirinus (Candéze) comb. n.

Lacon glivinus Candéze, 1865 : II.
Adelocera glivinus (Candéze) Fleutiaux, 1926 : 98.

LECTOTYPE (present designation). Fiji Is.: Ins. Fidgii; Janson coll. ex
Candéze, 1903.130; Lacon glirinus Cdz. Fidjee, Thorey [Cand.]; Lacon glirinus
Cdze., Type (ex coll. Cand.) [Janson] (BMNH). The abdomen is missing.

Agrypnus goudotii (Candéze) comb. n.

Lacon goudotii Candéze, 1857 : 106, pl. 2, fig. 1.
Lacon goudotii (Candéze) Fleutiaux, 1926 : 99.

Candéze’s comment that the last abdominal segment is sometimes smooth suggests
that he had specimens of both sexes before him at the time of the description.
He records the length and width as 25 mill. and 8} mill. respectively (I mm longer
than the figure). Up to the present time I have been unable to locate a specimen
of this size in the BMNH collections. The female from the Dejean collection
bearing Jansons determination label with the word ‘Type’ measures only 22 mm.

160 Cc. M. F. von HAYEK

Designation of the lectotype is postponed until a further search has been made
for Candéze’s material.

The generic attribution is based on the following specimen: 9, MADAGASCAR:
Janson coll. ex. Candéze, 1903.130; Lacon goudotii Cdz. Mad. [Cand.] (BMNH).

Agrypnus gracilentus (Schwarz) comb. n.

Lacon gracilentus Schwarz, 1902f : 306.
Adelocera gracilentus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected in the
Salomo Is. (Tulagi) by R. v. Bennigsen.

Syntype-material: ?DEI, Eberswalde.

Generic attribution based on the following specimens: SOLOMON Is.: I ex., Salomon
Is. Guadalcanal, Lavoro, H. T. Pagden, 16.11.34, Wild fig; Adelocera gracilenta
(Schw.) [Van Z.] (BMNH).

Agrypnus gracilis (Candéze) comb. n.

Lacon gracilis Candéze, 1874 : 83.
A deloceva (Compsolacon) gracilis (Candéze); Van Zwaluwenburg, 1966 : 289.

The description is based on an unrecorded number of specimens from ‘Gilolo,
Batchian, Aru, Waigioe, Kai, Mysol, Nlle Guinée.’ etc. Although this species formed
part of Candéze’s second collection (see p. 271), there can be no doubt that the
specimens listed below are syntypes, presumably acquired by Janson. The rest
of the material, which I have not seen, should be in the IRSNB, Brussels.

Syntypes examined. INDONESIA: I 9, Wagiou; Janson coll., ex Candéze, 1903.130;
Lacon gracilis Cdz., n.sp. Waigioe [Cand.]; Lacon gracilis Cand., Type ex coll.
Cand. [Gahan]. 1 ex., Wag. Wallace; ex coll. Saunders; Janson coll. 1903.130; —
gracilis Cdz. [Cand.]. 1 g, C. W. Gilolo; Janson coll. 1903.130; gracilis Cdz. [two
Candéze determination labels]. 1 g, C. W. Gilolo; Janson coll. 1903.130. I ex. (badly
damaged) Mysol; Janson coll. 1903.130; gracilis Cdz. [Cand.]. 1 9, Ins Kaisa (sic);
Janson coll. 1903.130; gracilis [Cand.]. 1 9, Ins Aru, Dept.; Janson coll. 1903.130.
24, N. W. N. Guinea/C. W.; Janson coll. 1903.130 (BMNH).

Agrypnus gragetensis (Szombathy) comb. n.

Lacon gragetensis Szombathy, 1909 : 119; fig. 2.
A delocera gragetensis (Szombathy) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). NEw GuINEA: 9, Ins. Graget, Biro; Lacon
gragetensis n.sp. [Szomb.] (TM, Budapest).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 161

Agrypnus granulatus (MacLeay) comb. n.

Lacon granulatus MacLeay, 1872 : 251.
Lacon granulatus (MacLeay); Neboiss, 1956 : 8.

The description is based on an unrecorded number of specimens from Gayndah
(Australia, Queensland) collected by Mr Masters. Neboiss states that the type is
in the AM, Sydney, but since he publishes no details concerning the specimen, his
statement cannot be accepted as a lectotype designation.

The generic attribution is based on the following specimen: AUSTRALIA: 4,
Gayndah, Queensland, Masters; Lacon granulatus McLeay [? MacLeay] (MNHN,
Paris). This specimen is almost certainly a syntype, presumably acquired by the
MNHN, Paris as a gift or by exchange.

Agrypnus griseopilosa (Fleutiaux) comb. n.
Adeloceva (Archontas) griseopilosa Fleutiaux, 19324 : 49.

LECTOTYPE (present designation). MaApacascar: 9, Diego Suarez; Type;
griseopilosa Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 29,1, same locality but without determination labels (MNHN,
Paris).

The specimens from the second recorded locality, Monts de Francais, [S. E. of
Diego-Suarez] have not been located.

Agrypnus grisescens (Candéze) comb. n.

Lacon grisescens Candéze, 1874 : 52.
Adelocera grisescens (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MADAGASCAR: 3, Madagas.; Janson coll.
ex Candéze, 1903.130; grisescens Cdz., type unique [Cand.]; Lacon grisescens
Cand. type [Gahan] (BMNH).

As the description was published after 1869 (see p. 271) one would expect to
find the type-material in the IRSNB, Brussels. However the only specimen
in that collection with Candéze’s determination label was collected on Nossi Be
by Brandt and measures 14mm, compared with the published length of 12 mm.
How this specimen came into Janson’s possession remains a mystery.

Agrypnus guttatus (Candéze) comb. n.

Lacon guttatus Candéze, 1857 : 151.
Lacon guttatus Candéze; Neboiss, 1956 : 8.

Holotype. AUSTRALIA: 3, Melb. B.; 63.10 [BMNH registration number =
G. R. Waterhouse collection, purchased of Stevens}; Lacon guttatus Cand. [Cand.];

Lacon guttatus Candéze, Type [C. O. Waterhouse] (BMNH).
i;

162 Cc. M. F. von HAYEK

I have not had the opportunity of examining the type of L. maculatus Macleay,
1872, which according to Lea (1920 : 379) is a small specimen of guttatus (Candéze).

Agrypnus gypsatus (Candéze) comb. n.

Lacon gypsatus Candéze, 18914 : 773.
Archontas gypsatus (Candéze) Fleutiaux, 1947 : 283.

The description is based on a large, but unrecorded number of specimens
collected by Fea in the Carin Cheba mountains in December 1888.

Syntype-material: the main part should be in the MCSN, Genoa, but various
workers appear to have obtained syntypes (see below).

Syntypes examined: BurMA: I g, Carin Cheba, goo-I000m. L. Fea v.xii-88;
co-type [BMNH curatorial label]; Lacon grypratus [sic] Cand.; Andrewes Bequest,
B.M. 1922: 221 (BMNH). 1 ex., same locality and date; L. gypsatus ex. type
Cdz. [? Cand.], Cand. Ann. Mus. Gen. 1888 et 1891 sec Gestro [Fleut.]. 1 ¢, Carin
Cheba, goo-1100m. L. Fea xii-88; L. gypsatus Cdz. n.sp. [Cand.]. 1 9, same
locality without determination label (MNHN, Paris). Candéze did not record
the species in 1888.

Agrypnus haedulus (Candéze) comb. n.

Lacon haedulus Candéze, 1857 : 122; pl. 2, fig. Io.
Adeloceva haedulus (Candéze) Fleutiaux, 1926 : 96.

Holotype. <3, Laf. Ind. bor. Bacon [Janson]; Janson coll. 1903 : 130; haedulus
Cdz., type [Cand.]; Lacon haedulus Cdze. Cand. Type e coll. de Laferté [Janson]
(BMNH).

The published locality is ‘Indes-Orientales boreales’, see p. 271.

Agrypnus hastatus (Candéze) comb. n.

Tilotarsus [sic] hastatus Candéze, 1857 : 177.
Centrostethus hastatus (Candéze) Schwarz, 1898c : 414.

The description is based on Madagascan specimens received by the author from
Klug as hastatus and M. de la Ferté Sénectére as ‘A. spimicollis in coll. Dejean’.

LECTOTYPE (present designation). MADAGASCAR: 4, Madagascar, Goud,
No. 16088 (MNHU, Berlin).

Paralectotypes. I 9, 16088; Madagascar Goud. (MNHU, Berlin). 1 4g,
Madagascar; Janson coll. 1903.130; Tilotarsus hastatus Cdze, type, Ag. spinicollis
Dej. ex. coll. Dejean [Janson]. 1 9, 1 g same data, but without the word ‘type’
on the determination label. I g, Madagascar, Janson coll. 1903.130; Tilotarsus
hastatus Cdz., Madag. [Cand.]; Tilotarsus hastatus Cdze, ex. coll. Cand. [Janson]
(BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 163

The MNHU, Berlin specimens stand beside Gerstaecker’s label: hastatus Cand.,
Madag. Goud. Despite the absence of Klug’s and Candéze’s labels, which is
probably due to Gerstaecker (see p. 275), these are believed to be the specimens
submitted, with the name hastatus, to Candéze by Klug. The number 16088 on
these specimens refers to a catalogue and list of localities in the MNHU, Berlin.

The three specimens from the Dejean collection are those submitted by de la
Ferté Sénectére. They are recorded in de la Ferté Sénectére’s manuscript list
(in BMNH) of Dejean specimens sent to Candéze for determination, The absence
of Dejean’s and/or Candéze’s determination labels and of any indication that they
formed part of de la Ferté Sénectére’s collection is probably due to Janson (see
p. 276). I have assumed that the specimen from the Candéze collection is one
retained by Candéze at the time of the description.

Agrypnus haterumana (Ohira) comb. n.
Colaulon (Cryptolacon) haterumana Ohira, 1967b : 104.
Holotype. Hateruma-jima near Ishigaki, 27.vii.1964 (M. Yasui). Ohira
collection. Not examined.
The generic attribution is based on the description.

Agrypnus herzi (Koenig) comb. n.

Lacon herzi Koenig, 1887 : 353.
Adelocera herzi (Koenig) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected in
Korea by O. Herz and submitted to Koenig by D. G. Sievers.

Type-material: ?ZI, Leningrad (Horn and Kahle, 1935 : 138).

The generic attribution is based on the description. The species bears a

resemblance to A. judex (Candéze) and A. setiger (Bates) in that erect setae are
present on the elytra.

Agrypnus hexagonus (Candéze) comb. n.
Tilotarsus [sic] hexagonus Candéze, 18936 : 13.
Lobotarsus hexagonus (Candéze) Schwarz, 1906 : 30.
The description is based on an unrecorded number of specimens from Antana-
narivo [MADAGASCAR].
Type-material: >IRSNB, Brussels.
The generic attribution is based on the following specimen: 2, Museum Paris,

Madagascar, Coll. Ch. Alluaud, 1904; Tilotarsus hexagonus Cand. [Cand.] Cand.
det. 1895 (MNHN, Paris).

164 Cc. M. EF: von HAYEK

Agrypnus hispidulus (Candéze) comb. n.

Lacon hispidulus Candéze, 1857 : 126.
Adelocera hispidulus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). JAva: 9, Dej. Java [Janson]; Janson
coll. 1903.130; Lacon hispidulus Cdze., Cand. Type e coll. Dejean [Janson]
(BMNH). The absence of Dejean and Candéze determination labels may be
ascribed to Janson (see p. 276).

Agrypnus holosericeus (Candéze) comb. n.

Lacon holosericeus Candéze, 1893 : 170.
Adelocera holosericeus (Candéze) Fleutiaux, 1926 : 96.

The description is based on a large number of specimens, collected by Bell in
Kanara, in the Andrewes collection. Since Candéze may have retained specimens
for his own collection, the designation of a lectotype is postponed until any
material in the IRSNB, Brussels has been examined.

Syntypes examined: Inp1A: Mysore, 1 g, Kanara; Type (BMNH curatorial
label]; Lacon holosericeus Cand. Kanara [Gahan]; Lacon holosericeus Cand. n.sp.
3 [?Candéze]. 1 g, Kanara; cotype [BMNH curatorial label]; Lacon holosericeus
Cand. 3 g, Kanara; Andrewes coll. BM. 1933.109. One specimen bears Andrewes
determination label: Lacon holosericeus Cand. 6 3, 1 9, Cotype [BMNH curatorial
label] Kanara; Andrewes Bequest BM, 1922.221. One specimen bears Andrewes
determination label: Lacon holosericeus Cand. (BMNH). 1 ex., Kanara; L.
holosericeus [? Cand.]; L. holosericeus Cand. Ann. Soc. Ent. Belge. 1893 : 170
(NMHN, Paris).

Agrypnus horni (Schwarz) comb. n.

Lacon lineatus Schwarz, 1901 : 18. [Homonym of Lacon lineatus Candéze 1896 : 8.]
Lacon horni Schwarz, 1902e : 155. [Replacement name for Lacon lineatus Schwarz. |
Adeloceva horni (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from CEYLON:
Matala, Kandy, Paradna.

Type-material: ?DEI, Eberswalde.

The generic attribution is based on the following specimen: J, Ceylon, Nietn.;
16119; Lacon lineatus Schw. [Schwarz] (MNHU, Berlin).

Agrypnus hottentota (Candéze) comb. n.

Lacon hotientota Candéze, 1857 : 138.
Adelocera hottentota (Candéze) Fleutiaux, 1926 : 96.

Holotype. SoutH Arrica: Laf. P.B.S. [Promontorium Bonae Spei]; Janson

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 165

coll. 1903 : 130; L. hottentota Cdz. Type [Cand.]; 276; Lacon hottentota Cdze.
Cand. Type coll. de Laferté [Janson] (BMNH).

The metasternum and wings are much reduced in length.

Agrypnus hova (Fleutiaux) comb. n.
Lacon scutellatus Fairmaire, 1903 : 202. [Junior primary homonym of Lacon scutellatus

Candéze, 1893.]
Adeloceva hova Fleutiaux, 1926: 99. [Replacement name for Lacon scutellatus Fairmaire. |

LECTOTYPE (present designation). MADAGASCAR: 3, Madag. Perrier [Fairm.];
Museum Paris, coll. Perrier de la Bathie 1906; type; Lacon [Fleut.] scutellatus
[Fairm.] (MNHN, Paris).

Paralectotypes. 1 ex., Madag. Perrier [Fairm.]; Museum Paris, coll. Perrier
de la Bathie 1906; scutellata Fairm. ex typ. [Fleut.]; hova Fleut., scutellatus Fairm.
nec Cand. [Fleut.] 1 ex., locality and collection labels as lectotype. 4 ex., locality
label as lectotype. The last five paralectotypes lack determination labels.
(MNHN, Paris).

The published locality is: Environs de Suberbieville (H. Perrier).

Agrypnus humilis (Erichson) comb. n.

Lacon humilis Erichson, 1842 : 136.
Lacon humilis Erichson; Neboiss, 1956 : 9.

LECTOTYPE (present designation). TASMANIA: 3g, Terra Van Diem. Schayer,
Nr. 16139 [museum catalogue number, refers to locality on label]; humilis Er. +
Cand. Van Dem. Schayer [Gerstaecker, mauve label] (MNHU, Berlin).

The description is based on an unrecorded number of specimens collected by
Schayer at Woolnorth [40°39’S., 144°43’E.], Tasmania and sent to the MNHU,
Berlin (Germar, 1842: 84). Candéze (1857: 184) states that he saw the single
specimen of this species in that museum. The absence of Erichson’s label is
probably due to Gerstaecker (see p. 275).

Agrypnus hypnicola (Kishii) comb. n.
Colaulon (Cryptolacon) hypnicola Kishii, 1964 : 33; pl. 1, figs 5, 6; pl. 3, fig. 9.

The description is based on a holotype, allotype and 311 paratypes from six
localities in JAPAN. Two paratypes are in the T. Shubata collection, but the
location of the remainder is not recorded. They are probably in the author’s
collection, and that of the Heinan High School, Kyoto. As the author records
the data on each specimen there should be no difficulty in recognizing the material.

The generic attribution is based on the description.

166 C. M. Fe von® HAYEK

Agrypnus illimis (Horn) comb. n.

Lacon illimis Horn, 1894 : 336.
Colaulon illimis (Horn) Arnett, 1952 : II9.

LECTOTYPE (present designation). U.S.A.: 9, Tucson, Arizona, Wickham;
396; Type No. 3329; L. illimis Horn [Horn] (ANS, Philadelphia). Length 8-5 mm.
The description is based on an unrecorded number of specimens from Tucson

(Wickham) and near Yuma, Arizona. Up to the present time only the specimen
designated as the lectotype has been located.

Agrypnus impressicollis (Elston) comb. n.
Lacon impressicollis Elston, 1924 : 202.
Lacon impressicollis Elston; Neboiss, 1956 : 9.
The description is based on an unrecorded number of specimens from TASMANIA,
George Town, Ben Lomond (4,000 ft).
Syntypes (see p. 274): SAM, Adelaide, AM, Sydney, NMU, Melbourne.

The generic attribution based on the following syntypes: 2 3, Tasmania, George
Town; co-type (SAM, Adelaide).

Agrypnus impressus (Candéze) comb. n.
Lacon impressus Candéze, 1878a : 102.
Adelocera impressus (Candéze) Fleutiaux, 1926 : 96.
Lectotype (designated by Van Zwaluwenburg, 1959 : 353). NEW GUINEA:
Andai, New Guinea (Beccari) (MCSN, Genoa). Not examined.

Paralectotype. I ex., recorded by Van Zwaluwenburg (loc. cit.) (MCSN, Genoa).
Not examined. Van Zwaluwenburg does not record any details of the determina-
tion labels on the specimens.

The generic attribution is based upon the description.

Agrypnus inaequalis (Fleutiaux) comb. n.

Adeloceva (Archontas) inaequalis Fleutiaux, 1934) : 52.

LECTOTYPE (present designation). MADAGASCAR: ?9, Madagascar, Baie
d’Antongil, A. Mocquerys; inaequalis Fleut., Type [Fleut.] (MNHN, Paris).

Paralectotypes. I g, 3 9, same locality as the lectotype without determination
labels. x 4, 2 9, Fianarantsoa, without identification labels. 1 9, Museum Paris,
Madagascar. Prov. de Fenerive, Reg. de Soanierana, A. Mathieux 1905, without
identification labels (MNHN, Paris).

rf

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 167

Agrypnus incertus (Fleutiaux) comb. n.
Adelocera incerta Fleutiaux, 1927 : 75.

LECTOTYPE (present designation). SoutH VIETNAM: 9, Cochinchine; voisin
de incertus, Cand. vidit [Fleut.]; pres longus et hispidulus [Fleut.]; Adelocera
incerta Fleut., Type [Fleut.] (MNHN, Paris).

Agrypnus incurvatus (Fleutiaux) comb. n.
Adelocera incurvata Fleutiaux, 1927: 81.

LECTOTYPE (present designation). NortH VIETNAM: 9, Hoa Binh, Tonkin;
Hoa Binh, Tonkin, de Cooman; 11; Adelocera incurvata Fleut., Type [Fleut.]
(MNHN, Paris).

Paralectotypes. 1 9, same locality as the lectotype but without a determination
label. 6 ex., Tonkin, Lac Tho, Hoa Binh, A de Cooman, one specimen bears
Fleutiaux’s determination label (MNHN, Paris).

Agrypnus indosinensis (Fleutiaux) comb. n.

Adelocera (Compsolacon) indosinensis Fleutiaux, 1927 : 86.
Compsolacon indosinensis (Fleutiaux) Fleutiaux, 1947 : 286.

LECTOTYPE (present designation). Laos: 9, Muong Nga, Luang Prab.
13.3.18; Laos, Luang Prabang (environs), Vitalis de Salvaza; Compsolacon indo-
sinensis Fleut. Type [Fleut.] (MNHN, Paris).

Paralectotype. NortH VIETNAM: I ex., Env. de Lam (Tonkin) Blaise;
indosinensis Fleut. [Fleut.}] (MNHN, Paris).

Agrypnus inductus (Candéze) comb. n.

Lacon inductus Candéze, 1888 : 668.
Adelocera inductus (Candéze) Fleutiaux, 1926 : 96.

The description is based on a single specimen from Bhamo, Haut Birmanie
[Burma] collected in June by Fea.
Holotype: MCSN, Genoa.

The generic attribution is based on the description.

Agrypnus indutissimus (Candéze) comb. n.

Lacon indutissimus Candéze, 1892a : 797.
Adelocera indutissimus (Candéze) Fleutiaux, 1926 : 96.

The description is based on a single specimen from Kifa-juc [ENGGANO I.]
collected by Modigliani in May—June 1891 and submitted to Candéze by Gestro.

Holotype: ?MCSN, Genoa.

168 Cc. M. F. von HAYEK

The generic attribution is based on Candéze’s comment that this species resembles
feralis Candéze (see p. 154).

Agrypnus inflatus (Candéze) comb. n.

Lacon inflatus Candéze, 1897 : 9.
Adelocera inflatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Mysore
[Inp1A].

Type-material: ?>IRSNB, Brussels.

The generic attribution is based on a specimen, which may be part of the syntype-
series; INDIA: I 9, Inde; Lacon inflatus Cand., Mysore [Cand.] Candéze det. [Fleut.]
(MNHN, Paris).

Agrypnus inops (Candéze) comb. n.

Lacon inops Candéze, 1874 : 67.
Adelocera inops (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from ‘Malacca,
Siam and Borneo’. The IRSNB, Brussels does not contain any specimens from
these localities (see p. 271). The only specimens under this name, a female
from Cambodia with Candéze’s yellow-bordered determination label and a male
from the same locality, are not conspecific with the lectotype and belong to a
species unknown to me.

LECTOTYPE (present designation). INp1A: 9, India [Janson]; L. inops Cdz.,
type [Cand.]; Lacon inops Cand. type, India [Gahan]; Janson coll. ex Candéze,
1903.130 (BMNH). The discrepancy between the published locality and the label
is probably due to Janson (see p. 276).

Paralectotype. BorRNEO: 9, Schwanez, Borneo; L. inops Cdz. type [Cand.]
(RNH, Leiden).

Agrypnus insignitus (Candéze) comb. n.

Lacon insignitus Candéze, 1874 : 98.
Lacon insignitus Candéze; Neboiss, 1956 : 9.

LECTOTYPE (present designation). AUSTRALIA: ¢, N. Holl. Q’land; L.
insignitus [Janson] Cand. (Janson ms.) [Gahan]; Lacon insignitus Cand. type,
Queensland [Gahan] (BMNH).

Paralectotype. 2 ¢, N. Holl. Q’land.; Janson coll. 1903 : 130; Lacon insignitus
Cand. [Gahan]. One specimen bears Candéze’s label: L. insignitus n.sp. (BMNH).

The wings and metasternum are much reduced in length.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 169

Agrypnus insularis (Candéze) comb. n.

Lacon insularis Candéze, 1874 : 74.
Adelocera insularis (Candéze) Fleutiaux, 1926 : 98.

The description is based on an unrecorded number of specimens of variable
appearance from the Indian Archipelago, the Moluccas, New Guinea, Batchian
and Malacca. Candéze comments that it was found often by Wallace.

Lectotype (designated by Van Zwaluwenburg, 1959 : 353). INDONESIA: 4g,
C. W. Mysol; Janson coll. 1903.130; insularis [Cand.] (BMNH). In view of the
fact that Candéze’s second collection (see p. 171) is in the IRSNB, Brussels it is
strange that Van Zwaluwenburg selected the specimen in the BMNH as the
lectotype. However as it agrees with the description, comes from the published
locality and bears Candéze’s determination label, there is no justification for
rejecting Van Zwaluwenburg’s action.

Paralectotypes. Van Zwaluwenburg records that the lectotype is the first of
nine specimens but he does not indicate whether he believes the remaining eight
specimens to be part of the original syntype series. I have been able to discover
only the following seven specimens from the type-localities in the BMNH. I ex.,
only right elytron remaining, labels as lectotype. 2 3g, N.W. New Guinea/C.W.;
Janson coll. 1903.130; insularis [Cand.] 2 9, same locality and collection, one bears
a note ‘pas de sillons’ in Candéze’s hand writing. 9, Batchian/C.W. gl.; Janson
coll. 1903.130; Lacon insularis Cand., named by Candéze [Gahan]. This specimen
differs from the New Guinea females, which I believe to be correctly identified,
in that there is no pore on the fourth visible abdominal sternite. It belongs to
a species unknown to me. I 9, Waigou/C.W.; Janson coll. 1903.130; Lacon
insularis Cand., named by Candéze [Gahan]. This specimen also differs from
the New Guinea specimens in that there is a pore on both the third and fourth
visible abdominal sternite. It belongs to a species unknown to me (BMNH).
I 9, Malacca [sic]; Lacon insularis Cdz. Molucc. [sic] [Candéze, yellow border];
Collection E. Candéze; Lacon insularis Cd. det. E. Candéze [IRSNB, Brussels
curatorial label] (IRSNB, Brussels). The different localities on the two labels
are probably due to an error in copying. The specimen measures 7:5 mm, compared
with a published length of 5-6mm. The fifth visible abdominal sternite bears
a shiny patch and there is no pore on the fourth sternite. The specimen is not
conspecific with the paralectotype from New Guinea in the BMNH. 1 3, Malacca
Collection E. Candéze; Lacon insularis Cd. det. E. Candéze [IRSNB, Brussels
curatorial label] (IRSNB), correctly identified.

The specimen (2) from Sarawak [Borneo] (BMNH) recorded by Van Zwaluwenburg
(loc. cit.) is correctly identified.

Agrypnus insulsus (Candéze) comb. n.

Tilotarsus [sic] insulsus Candéze, 1895) : 57.
Lobotarsus insulsus (Candéze) Schwarz, 1906 : 30.

170 Co MM. Bo von TAY EK

The description is based on an unrecorded number of specimens collected at
Diego Suarez [MADAGASCAR] by Ch. Alluaud.

Type-material: P>MNHN, Paris and/or IRSNB, Brussels.

The generic attribution is based on the following specimen which may be a
syntype; J, Museum Paris, Madagascar, Coll. Ch. Alluaud 1904; Tilotarsus insulsus
Cand. Cand. det. 1895 (MNHN, Paris).

Agrypnus intermedius (Schwarz) comb. n.

Lacon intermedius Schwarz, 1902b : 199.
Adelocera intermedius (Schwarz) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Borneo: g, N. Borneo; coll. Schwarz;
Typus; intermedius Schw. [Schw.] (DEI, Eberswalde).

Paralectotypes. 1 ¢, Borneo, Kina Balu; coll. Schwarz; Typus 2, N. Borneo;
coll. Schwarz; Typus (DEI, Eberswalde). 1 g, Kina Balu; coll. Schwarz Typus.
3, Borneo, Kina-Balu; Coll. Schwarz; Typus (DEI, Eberswalde). These two
specimens are distinct species, which are not conspecific with the lectotype. They
belong to species unknown to me.

Agrypnus interpunctatus (Klug) comb. n.

Elater (Agrypnus) interpunctatus Klug, 1833 : 66 (1834 : 154).
A delocera interpunctatus (Klug) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MADAGASCAR: g, 16104 [museum cata-
logue number = information on next label]; Madagasc. Goud. (MNHU, Berlin).

Paralectotype. 3, Madagasc. Goud. Nr. 16104 (NMHU, Berlin). The specimens
stand beside a blue label in Gerstaecker’s handwriting: interpunctatus Cand.
Elater interpunctatus Klug+ Madag. Goud. The absence of Klug’s determination
label is probably due to Gerstaecker (see p. 275).

Agrypnus irroratus (Klug) comb. n.
Elater (Agrypnus) tvroratus Klug, 1833 : 65 (1834 : 153).
Lacon irroratus (Klug) Germar, 1840 : 265.

Lacon confusus Candéze, 1857 : 109, pars. [Synonymized by Candéze, 1874 : 53.]
A delocera ivrovatus (Klug) Fleutiaux, 1826 : 99.

Elater (Agrypnus) irroratus Klug. LECTOTYPE (present designation). MapaA-
GASCAR: 3, Madag. Goudot, No. 16096 [museum catalogue number = Madagascar,
collected by Goudot] (NMHU, Berlin).

Paralectotypes. 2 9, 16096; Madag. Goudot, No. 16096. 1 9, 16096; Madag.
Goudot (MNHU, Berlin). The specimens stand beside a blue Gerstaecker label:
irroratus Cand., Elat. irroratus Klug, Madagascar, Goudot. The absence of Klug’s
determination label is probably due to Gerstaecker (see p. 275).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 171

Lacon confusus Candéze. The description is based on an unrecorded number
of specimens from Madagascar received as micans from Klug and binotatus from
Chevrolat. The length is recorded as 12-24 mm, but, as the corresponding widths
are 3+-42mm, it seems probable that the second measurement is a misprint,
and should read 14mm. Candéze (1874 : 53) synonymized confusus with Lacon
wrroratus Klug, which he believed to be very variable in size. Fleutiaux (1899 : 222)
states that Candéze’s confusus material consisted of two ‘formes’ (species in this
context) and that the type of the larger one was in the MNHU, Berlin. I have
not been able to find this specimen in the MNHU, Berlin collections. The BMNH
contains a female bearing the following labels: Madagascar; Janson coll. ex.
Candéze, 1903.130; Lacon confusus Cdz. Mad. Mnz. [Cand.]. The specimen is
I4°5mm long, agrees well with Candéze’s description and is conspecific with
the lectotype of ivrvoratus Klug. If further searches fail to discover Klug’s ‘micans’
specimen, I suggest that this specimen be designated as the neotype of Lacon
confusus Candéze.

The specimen which Fleutiaux regarded as the type of Candéze’s smaller species
is a male in his own collection (now in MNHN, Paris) bearing the following labels:
L. confusus type, Mon. 1, p. 109, Fleut. Bull. Mus. 99 : 22 [Fleut.]. The specimen
stands beside a Chevrolat collection label: Lacon confusus Cand. Mon. 1, 1857,
p. 109.16. Madagascar. The specimen measures I1°5mm and agrees with
Candéze’s description. Despite the fact that it lacks any kind of label indicating
that it was Chevrolat’s binotatus, I believe that it formed part of Candéze’s original
confusus material. It is not conspecific with ivoratus Klug and belongs to a
species unknown to me.

GERMAR’S INTERPRETATION OF Lacon irroratus (KLUG). Germar (1840 : 265)
refers to Klug’s description of zvroratus but he does not record whether he saw
Klug’s material. The BMNH contains a male with the following labels: irroratus
Kl. Madag. [Germar]; Janson coll. ex. Schaum, 1903.130; Md. [mss. on triangular
blue label]; 1015. I believe that this specimen, which is conspecific with the
lectotype of zvroratus Klug, is that on which Germar based his interpretation of
the species.

Agrypnus jansoni (Fairmaire) comb n.

Lacon jansoni Fairmaire, 1871 : 40.
Adeloceva jansoni (Fairmaire) Fleutiaux, 1926 : 96.

The description was based on two specimens, one from Bourbon [REUNION]
in the de la Ferté collection, which Fairmaire states Candéze believed to be a
variety of L. irroratus (as virroratus, a misprint) and a second specimen in the
same collection labelled Madagascar in Coquerel’s hand. I have not been able

_ to find these specimens, which should be in the BMNH. If Fairmaire acquired

these specimens they may now be in the MNHN, Paris or IRSNB, Brussels,
Bee p. 275.
The generic attribution is based on the following specimen: 1 9, Madagascar.

172 C. M. BY von THAWEK

Diego Suarez 3, Ch. Alluaud, 1893; L. jansoni Fairm. [Cand.] Cand. det. 1895
[Fleut.] (MNHN, Paris).

Agrypnus judex (Candéze) comb. n.

Lacon judex Candéze, 1874 : 62.
A delocera judex (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). CHINA: 9, Shanghai [Janson]; Janson
coll. 1903 : 130; L. judex Cdz. [Cand.]; L. judex Cand. Type [Gahan] (BMNH).

Agrypnus jurulosus (Candéze) sp. rev., comb. n.

Lacon jurulosus Candéze, 1889 : 73.

The description is based on an unrecorded number of specimens from SIKKIM,
Himalaya. I have been unable to find this material in the IRSNB, Brussels or
elsewhere.

Schenkling (1925 : 27) records this species as a synonym of truncatus Herbst,
but without giving any reason for doing so. Since Candéze implies that the
species bears a resemblance to L. coctus Candéze, I believe Schenkling’s synonymy
to be erroneous.

The generic attribution based on the following specimen; INnp1A: 9, Kullu,
1019.6; jurulosus Cand. Beng. Barwai C. [Candéze, yellow border]; Collection
E. Candéze; jurulosus Cd. det. E. Candéze [IRSNB curatorial label] (IRSNB,
Brussels). This may be one of the large number of specimens recorded by
Candéze (1892 : 484). It is not conspecific with truncatus Herbst.

Agrypnus kenyensis (Fleutiaux) comb. n.

Lacon kenyensis Fleutiaux, 1919 : 18.
Adelocera kenyensis (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). KrEnya: g, Afr. or. Anglaise, Mt. Kenya
vers ouest, Zone de Foréts, Alluaud & Jeannel; Forets inferieure, Podocarpus,
2400m, Jan., Fevr. 1912, st. 39; Lacon kenyensis Fleut. type. Voy. All. Jeann.
Afr. or. Elat. p. [Fleut.] (MNHN, Paris).

Paralectotypes. g ex., same locality as the lectotype, 2 specimens bear
Fleutiaux’s determination label (MNHN, Paris). 1 ex., Afr. or. Anglaise, Mt.
Kenya vers ouest. Zone de foréts, Alluaud & Jeannel; Praires découvertes entre
les rivieres Amboni et Narémuru (Abt. 1800-2000 m) jan—fev. 1912, st. 30 et 50;
Adelocera (Lacon) kenyensis Fleut. [Fleut.] (BMNH). 7 ex. Museum Paris, Afr.
Orient. Angl. Escarpment (Wa Kikou-you) [Kikuyu in description] Ch. Alluaud.
One specimen bears the date ‘Aout’ and two others Fleutiaux’s determination
label (MNHN, Paris). Fleutiaux recorded 8 specimens from Kikuyu, but only
7 examples have been located.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 173

Agrypnus kinangopa (Fleutiaux) comb. n.

Adelocera kinangopa Fleutiaux, 1935e@ : 92.

LECTOTYPE (present designation). KENyA: 9, Turner, Kinangop, March, 30;
Museum Nairobi; Adelocera (Lacon) Kinangopa Fleut. Type [Fleut.] (MNHN,
Paris).

Paralectotypes. 2 ex., same locality as lectotype, with Fleutiaux’s determination
label (MNHN, Paris, NM, Nairobi).

Agrypnus labiosus (Candéze) comb. n.

Lacon labiosus Candéze, 1874 : 86.
Adelocera (Compsolacon) labiosus (Candéze) Van Zwaluwenburg, 1959 : 353.

Lectotype (designated by Van Zwaluwenburg, 1959 : 353). AUSTRALIA: d,
W. Australia/Fremantle [Janson]; Janson coll. 1903.130; Lacon labiosus Cand.
Type [Gahan]; Lacon sp. nov., impressions tarsals sur le pro et metathorax.
L. labiosus mihi. Champion Bay [28°44’S., 113°36’E.] and Fremantle, W. Australia
[Janson] (BMNH).

Paralectotype. 9, W. Australia/Fremantle; Janson coll. 1903.130; Lacon
labiosus Cand. cotype [Gahan] (BMNH).

The published locality is Swan Riv. Fremantle lies at the mouth of the Swan
River. The discrepancy between the published locality and the label is probably
due to Janson (see p. 276).

The description is misleading as it suggests that the tarsal grooves are deep
with well defined margins as in caliginosus. In fact, true grooves are absent,
their place being taken by depressions without well defined margins. The sides
and base of the depressions resemble the surface of the underside, being punctured
and clothed with scales.

Agrypnus lacrymosus (Candéze) comb. n.

Lacon lacrymosus Candéze, 1874 : 96.
Lacon lacrymosus Candéze; Neboiss, 1956 : 9.

LECTOTYPE (present designation). AUSTRALIA: 9, Janson coll. 1903.130;
lacrymosus Cdz. Type [Cand.]; Lacon lacrymosus Cand. S. Australia [Gahan]
(BMNH).

Paralectotype. 9, South Australia, C.W.; Janson coll. 1903.130; L. lacrymosus
Cdz. [Cand.] (BMNH).

The description is based on material in the Janson and Candéze collections.
The IRSNB, Brussels may possess additional paralectotype material.

Agrypnus lakhoni nom. n.

Adelocera (Archontas) subtuberculata Fleutiaux, 1927 : 83, pl. 1, fig. 2.

}
(
| Archontas subtuberculata (Fleutiaux) Fleutiaux, 1947 : 281. [Junior secondary homonym of

Agrypnus subtuberculatus (Schwarz, 1898a).]

174 CG. MM. E. von HAE K

LECTOTYPE (present designation). Laos: g, Mt. Laos; Adelocera subtuber-
culata Fleut. Type [Fleut.]; sp. ?>Ht. Laos, Vitalis; type de la fig. [Fleut.] (MNHN
Paris).

Paralectotypes. 2 g, 2 9, same locality as the lectotype. One specimen of
each sex bears Fleutiaux’s determination label (MNHN, Paris).

I have not seen the material from the second recorded locality, Lakhon (Harmand).

Agrypnus lameyi (Fleutiaux) comb. n.

Adeloceva (Compsolacon) lameyi Fleutiaux, 1927 : 85.
[Lacon muticus Herbst; Fleutiaux, 1918d : 194. Misidentification.]
Compsolacon lameyi (Fleutiaux) Fleutiaux, 1947 : 287.

LECTOTYPE (present designation). NorTH VIETNAM: 9, Hoa Binh, Cooman;
Compsolacon lameyi Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. I ex., Hagiang, Tonkin, Vitalis; Lacon muticus Hbst., Cand.,
Fl. 1918 [Fleut.]; Compsolacon lameyi Fleut., muticus Fleut, 1918 non Hbst.
[Fleut]. 1 ex. [illegible ? Ha-Lang] Lamey; Lacon musculus Cand.; Lacon muticus
Hbst., Cand. [Fleut.] (MNHN, Paris).

Agrypnus lapideus (Candéze) comb. n.

Lacon lapideus Candéze, 1857 : 141.
Compsolacon lapideus (Candéze) Fleutiaux, 1947 : 286.

LECTOTYPE (present designation). Inpb1A: g, India: Janson coll. 1903.130;
Lacon lapideus Cdze. Type e coll. Schaum [Janson]; lapideus [Cand., stuck to
underside of Janson label]; India or. [Cand., stuck to underside of Janson’s label]
(BMNH).

The published locality is ‘Indes Orientales’, see p. 271.

Agrypnus lateralis (Schwarz) comb. n.

Lacon lateralis Schwarz, 19030 : 381.
Adelocerva (Compsolacon) lateralis (Schwarz); Van Zwaluwenburg, 1966 : 208.

The description is based on an unrecorded number of specimens from ‘AUSTRALIA’
submitted to Plason by Vienna.

Type-material: ?DEI, Eberswalde.
The generic attribution is based on the description.

Agrypnus laticollis (Candéze) comb. n.

Lacon laticols Candéze, 1857 : 146; pl. 2, fig. 12.
Lacon laticollis (Candéze) ; Neboiss, 1956 : 9.

LECTOTYPE (present designation). AUSTRALIA: g, Australia; Janson coll.,

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 175

ex Candéze, 1903.130; Lacon laticollis Cdz. A. Thory [Cand.]; Lacon laticollis
Cand. Type ex coll. Candéze [Gahan] (BMNH).

Paralectotype. 9, N. Holl.; 18; laticollis Ch. v. Poll, L. laticollis [Cand.]:
Coll. Chevrolat: laticollis Cand., typ. mon. [Fleut.] (MNHN, Paris).

Agrypnus latiusculus (Candéze) comb. n.

Lacon latiusculus Candéze, 1878 : LIII (7).
Adelocera latiusculus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from
Madagascar.

Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: MADAGASCAR: Q,
latiuscula Cand., comparé au type, Fleut. [Fleut.] (MNHN, Paris).

Agrypnus latus (Candéze) comb. n.

Lacon latus Candéze, 1857 : 136.

Lacon flavipes Candéze, 1857 : 137; pl. 2, fig. 14. Syn.n.

Lacon maillardi Deyrolle, 1862 : 4; pl. 20, fig. 2. [Synonymized with flavipes Candéze, by
Candéze, 189Ic : 19.]

Adelocera latus (Candéze) Fleutiaux, 1926 : 96.

Lacon latus Candéze. LECTOTYPE (present designation). MADAGASCAR: g,
Madagascar; latus Cdz. Mad. Coll. Mn. [Cand.]; Lacon latus Cand., det. E. Candéze
[IRSNB curatorial label}; Collection Candéze (IRSNB, Brussels).

Paralectotype: 3, Dej. Madagascar; Janson coll. 1903.130; Lacon latus Cdze.
Cand. type [Janson]; Agrypnus gravis Dej. Cat. e coll. Dej. [Janson] latus [Cand.]
(BMNH).

Lacon flavipes Candéze. Holotype. ¢, Laf. Madagascar; Janson coll. 1903.130;
Lacon flavipes Cdze. Cand. Type e coll. de LaFerté [Janson]; L. flavipes voisin
de latus Cand. [Cand.] (BMNH). Candéze states that the specimen has lost all
the scales covering the body. Cleaning of the specimen has shown that it is
clothed with narrow scales, many of which have been lost.

Lacon maillardi Deyrolle. LECTOTYPE (present designation). REUNION: 9,
Lacon maillardi Deyr. Bourbon [believed to be Deyrolle’s handwriting]; Janson
coll. 1903.130 (BMNH).

Agrypnus lavaudeni (Fleutiaux) comb. n.
Adelocera (Lacon des Auteurs) lavaudeni Fleutiaux, 1932c : 153.
LECTOTYPE (present designation). MApaAGaAscaR: 9, Forét de Tempina, Est

Madagascar, sud Tamatave, Lavauden; Adelocera (Lacon) lavaudeni Fleut., type
[Fleut.] (MNHN, Paris).

176 CoM. 1B. Vion) TEU ASE

Agrypnus laxatus (Candéze) comb. n.

Lacon laxatus Candéze, 1895) : 56.
Adelocera laxatus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MApDAGAscAR: 4, Madagascar, Diego
Suarez, Ch. Alluaud 1893; Museum Paris, Madagascar, coll. Alluaud 1903; L. laxatus
n.sp. Cand. det. 1893 [Cand.] (MNHN, Paris).

Paralectotypes. I J, 3 ex., same locality and collection as the lectotype. All
bear Fleutiaux’s determination labels and one bears an additional label: Lacon
laxatus Cand. Ann. Belge 1859 p. 56 ex type [Fleut.] (MNHN, Paris).

The published locality is Montagne des Francais, 600-800 m.

Agrypnus leprosus (Candéze) comb. n.

Lacon leprosus Candéze, 1857 : 107; pl. 11, fig. 3.
Adelocera leprosus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens with a reference
to Agrypnus hydropictus Dup. in Dej. cat. ed. 3 : 99, and a specimen in the MNHU,
Berlin named leprosus by Klug. The designation of a lectotype is postponed
until any material extant in the MNHU, Berlin can be examined.

Syntype examined. MapaGascar: 4, Dej. Madagascar; Janson coll. 1903.130;
Lacon leprosus Cdze. [Janson]; Agrypnus hydropictus Dej. Cat. e coll. Dejean
[Janson]; L. leprosus [Cand.] (BMNH).

Agrypnus leucaspis (Candéze) comb. n.

Lacon leucaspis Candéze, 1874 : 63.
Adeloceva leucaspis (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected in
Malacca by Castelnau, in the Janson and Candéze collections. Selection of a
lectotype is postponed until any material extant in the Candéze collection (IRSNB,
Brussels) can be examined.

Syntypes examined. I ex., no locality label. Janson coll. 1903.130; L. leucaspis
Cdz. Type [Cand.]. The abdomen is missing. 1 9, Malacca; Janson coll. 1903.130;
L. leucaspis Cdz. [Cand.]. 1 9, 1 9, Malacca, Janson coll. 1903.130 (BMNH).

Agrypnus lezeleucii (Candéze) comb. n.

Lacon lezeleucir Candéze, 1857 : 158.
Colaulon lezeleuci (Candéze) Golbach, 1968 : 108.

LECTOTYPE (present designation). Mexico: 9, labelled as g, Laf. Vera Cruz,
Mex.; Janson coll. 1903.130; Lacon lezeleucii Cand. type, e coll. de Laferté [Janson]
(BMNH). The absence of Candéze’s determination label is probably due to Janson
(see p. 276).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 177

Paralectotype. d, labelled as 9, Laf. Vera Cruz, Mex.; Lacon lezeleucii Cand.
Cand. e coll. de Laferté [Janson]; Vera Cruz, lezeleucii [?Cand.] (BMNH). The
error in the determination of the sex is due to Champion (1894 : 263) (see p. 117).

Agrypnus limosus (Candéze) comb. n.

Lacon limosus Candéze, 1882 : Io.
Adeloceva limosus (Candéze) Fleutiaux, 1927 : 96.

The description is based on an unrecorded number of specimens from New
Guinea.
Type-material; ?>IRSNB, Brussels.

Generic attribution is based on the following specimen; NEw GuINEA: Papua;
Kokoda, 1200 ft, VII.33, determined by Van Zwaluwenburg (BMNH).

Agrypnus lindensis (Blackburn) comb. n.

Lacon lindensis Blackburn, 1892@ : 504.
Adeloceva lindensis (Candéze); Van Zwaluwenburg, 1959 : 352.

Lectotype (designated by Van Zwaluwenburg, 1959 : 352). @, 891, T [on card
mount, see p. 270]; Lacon lindensis Blackb. [Blackb.] (BMNH).

: Paralectotypes. An unknown number of cotypes in the SAM, Adelaide recorded
by Neboiss (1956 : 9).

| The published locality is ‘South Australia; not rare near Port Lincoln, under

_ bark of Eucalyptus’.

Agrypnus lolodorfensis nom. n.

Lobotarsus decoratus Schwarz, 1898a : 131.
Agrypnus decoratus (Schwarz) comb. n. [Junior secondary homonym of Agrypnus decoratus
(Candéze, 1881).]

The description is based on an unrecorded number of specimens collected in
Kamerun, Lolodorf by Conrad and submitted by Kraatz.

LECTOTYPE (present designation). CAMEROUN: 9, Kamerun; coll. Schwarz;
| histor Exempl. vielleicht Type [printed] decoratus Schw. [Schw.] (DEI, Eberswalde).

Agrypnus longicornis (Fleutiaux) comb. n.
| Adeloceva (Archontas) longicornis Fleutiaux, 1934) : 52.
| LECTOTYPE (present designation). MapaGascarR: 9, Manandriana, G.
Olsoufieff; Adelocera longicornis Fleut. Type [Fleut.] (MNHN, Paris).

Paralectotype. dg, Tananarive, G. Waterlot, 1919; Adelocera (Archontas)
Jongicornis Fleut. [Fleut.] (MNHN, Paris).
M

178 Cc. M. F. von HAYEK

Agrypnus longipennis (Schwarz) comb. n.

Lacon longipennis Schwarz, 1899 : 130.
Adelocera longipennis (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Madagascar.

Type-material: ?DEI, Eberswalde.

The generic attribution is based on the following specimen: 3, MADAGASCAR,
compared with the type by Fleutiaux (MNHN, Paris).

Agrypnus longus (Fleutiaux) comb. n.

Lacon longus Fleutiaux, 1903a@ : 571.
Archontas longus (Fleutiaux) Fleutiaux, 1947 : 281.

LECTOTYPE (present designation). SOUTH VIETNAM: 9, Cochin China; Lacon
longus Fleut. Type [Fleut.]; Lacon longus Fleut., type, Ann. Soc. ent. Fr. 1902 : 571
[Fleut.] (MNHN, Paris).

Agrypnus lopezi (Fleutiaux) comb. n.
Adeloceva (ex Lacon) lopezi Fleutiaux, 1934¢ : 477.

LECTOTYPE (present designation). PHILIPPINES: 3, La Carlotta, Occ. Negros
P.I.; vii.31; Al. Lopez, collector; F. C. Haddon collection; lopezi Fleut., type
[Fleut.] (MNHN, Paris).

Paralectotypes: I 9, same locality, collector and collection labels but without
a determination label (MNHN, Paris). An unknown number of specimens in
the HSPA, Honolulu.

Agrypnus luctus (Fleutiaux) comb. n.
Tylotarsus Lobitarsus (sic) luctus Fleutiaux, 1935¢ : 92.

LECTOTYPE (present designation). KEnya: 3, Lower Tana, Sabaki, avril-mai
(Turner & Macarthur); luctus Fleut. type [Fleut.] (MNHN, Paris).

Agrypnus lupinosus (Candéze) comb. n.

Lacon lupinosus Candéze, 1857 : 130.
Sagojo [sic] lupinosus (Candéze) Ohira, 1971¢ : 66.

Holotype. 3, Laf. Ind. Bor. Bacon [Janson]; Janson coll. 1903 : 130; lupinosus
Cdz. type [Cand.]; 47; Lacon lupinosus Cdze. Cand. Type e coll. de Laferté [Janson]
(BMNH).

The published locality is ‘Indes-Orientales boreales’, see p. 171.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 179

Agrypnus lustratus (Candéze) comb. n.

Lacon lustvatus Candéze, 1890 : cxlix.
Adelocera lustvatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected in
Chota Nagpore by Cardon (see p. 273).

Syntypes examined. INDIA: I ex., 13394 [Fry catalogue number = Lacon
lustratus Cand. determined by Cand., India or. Chota Nagpore, received from
Donkier, Dec. 1895]. Chota Nagpur; India orient.; Fry coll. 1905.100. 1 g, Chota
Nagpur, India orient; Fry coll. 1905: 100; Lacon lustratus (BMNH). 1 ex.,
lustratus n.sp. Cand. det. [Fleut.]; Chota Nagpore, Bengale; Cand. C. R. Soc.
Ent. Belge, 1890: 149, Ann. Soc. Ent. Belge, 1892 : 454. I ex, lustratus Cand.
Chota Nagpore, Bengale. 10 ex, Chota N. Bengale (MNHN, Paris). The selection
of a lectotype is postponed pending further study of the material.

Agrypnus lutosus (Candéze) comb. n.

Lacon lutosus Candéze, 1857 : 119.
Adelocera lutosus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from ‘des Indes
orientales’ (see p. 271) in the Schaum collection. As the species is not recorded
in the BMNH manuscript catalogue of the Schaum collection, it is assumed to
be lost. For the history of the Schaum collection see p. 280.

The generic attribution is based on the following specimen: NEPAL: g, Hardwicke,
Nepal; lutosus Cdz. [Cand.] (BMNH).

Agrypnus macgregori (Fleutiaux) comb. n.
Adeloceva (Archontas) macgregori Fleutiaux, 1934e : 364.

LECTOTYPE (present designation). PHILIPPINES: 9, ‘Culasi, Panay, 24 May’
18. P. I. R. C. McGregor; Adelocera macgregori Fleut. Type [Fleut.] (MNHN,
Paris).

Agrypnus macroderus (Candéze) comb. n.

Lacon macrodevus Candéze, 1865 : to.
Adeloceva macroderus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Batyan: g, Batchian; Janson coll. ex
Candéze 1903.130; Lacon macroderus Cdz. suppl. Batchian [Cand.]; Lacon

| Macroderus Cdze Type ex coll. Cand. [Janson] (BMNH).

Paralectotypes. 1 9, Batchian; Janson coll. 1903.130. 1 9, Batchian, C.W.;

| Janson coll. 1903.130. 1 ex, Bac.; L. macroderus Cdz. E. N.; Janson coll. 1903.130

(BMNH).

180 C. M. F. von HAYEK

Agrypnus maculosus (Macleay) sp. rev., comb. n.
Lacon maculatus Macleay, 1872 : 251.
The description is based on an unrecorded number of specimens collected at
Gayndah [AUSTRALIA, Queensland] by Masters.
Type-material: AM, Sydney, according to Neboiss (1956: 8).

According to Lea (1920 : 379) the type is a rather small specimen of L. guttatus
Candéze (see p. 161), a species described from Melbourne, Victoria. I have not
had the opportunity of examining Macleay’s material to confirm the synonymy.

The generic attribution is based on Lea’s proposed synonymy.

Agrypnus maculatus (Candéze) comb. n.

Lacon maculosus Candéze, 1874 : 53.
Lacon maculatus Brancsik, 1893 : 234. [Unjustified emendation. ]
Adelocera maculosus (Candéze) Fleutiaux, 1926 : 90.

LECTOTYPE (present designation). MaApaAGaAscar: 9, Laf. Madagascar; Mad.;
Coquerel; nov.sp. Madag.; Janson coll. 1903.130; Lacon maculosus Cand. type
[Gahan]; maculosus Cdz. type [Candéze] (BMNH).

Agrypnus madecassus (Fleutiaux) comb. n.
Adeloceva madecassa Fleutiaux, 1932é : 153.

LECTOTYPE (present designation). MapDAGaAscaR: 9, Forét de Tempina,
Est. Madag. Sud Tamatave, Lavauden; Adelocera madecassa Fleut., type [Fleut.]
(MNHN, Paris).

Agrypnus major (Candéze) comb. n.
Tilotarsus major Candéze, 1857 : 172.

Holotype. Mapacascar: g, Madagascar; Janson coll. 1903.130; major Cdze,
Type (e coll. de Mniszech) [Janson] (BMNH). The absence of a Candéze determi-
nation label is probably due to Janson (see p. 276).

Agrypnus mamillatus (Candéze) comb. n.

Lacon mamillatus Candéze, 1857 : 144; pl. 2, fig. 4.
Lacon mamullatus Candéze; Neboiss, 1956 : Io.

Holotype. AUSTRALIA: 9, Laf. Nov. Holl.; Janson coll. 1903.130; L. mamillatus
Cand. type [Cand.]; Lacon mamillatus Cdze Cand. type (e coll. de Laferté) { Janson]
(BMNH).

Agrypnus mansuetus (Blackburn) comb. n.

Lacon mansuetus Blackburn, 18926 : 288.
Adelocera (Compsolacon) mansueta (Blackburn); Van Zwaluwenburg, 1959 : 353.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 181

Lectotype (designated by Van Zwaluwenburg, 1959: 353, see p. 9).
AUSTRALIA: g, 4393, Narr. T. [on card mount, see p. 270]; Australia, Blackburn
Coll.; Lacon mansuetus Blackb. [Blackb.] (BMNH).

Paralectotypes. An unrecorded number of paratypes (cotypes) in SAM,
Adelaide, recorded by Neboiss (1961 : 6).

The published locality is N. S. Wales, taken by Mr Musson near Narrabri.
The wings and metasternum are much reduced in length.

Agrypnus marginatus (Candéze) comb. n.

Lacon marginatus Candéze, 1874 : 91.
Lacon marginatus Candéze; Neboiss, 1956 : Io.

LECTOTYPE (present designation). AUSTRALIA, N.S.W.: 3, Clarence River;
marginatus Cdz., Clar. Riv. [Candéze, red border]; Collection E. Candéze; Lacon
marginatus Cd. det. E. Candéze [IRSNB, curatorial label] (IRSNB, Brussels).

: Paralectotypes. AUSTRALIA, N.S.W.: 2 3, Clarence River; Collection E. Candéze;
: Lacon marginatus Cd., det. E. Candéze [IRSNB curatorial label] (IRSNB, Brussels).

The BMNH collection contains a pin with the following labels: Janson coll.
: 1903.130; Lacon marginatus Cand. Clarence River [?Gahan]; L. marginatus Cdz.,
| type [Cand.]. The specimen is missing and there are no additional specimens
| from the Janson collection from the type-locality which agree with the description.

Agrypnus marmoratus (Candéze) comb. n.

Lacon marmoratus Candéze, 1874 : 94.
Adelocera (Compsolacon) marmorata (Candéze); Van Zwaluwenburg, 1959 : 354.

Lectotype (designated by Van Zwaluwenburg, 1959: 354, see p. 9).
AUSTRALIA: 2, Janson coll. I903.130; marmoratus Cdz. type [Cand.]; Lacon
marmoratus Cand., Queensland [Gahan] (BMNH).

Since Candéze described this species after 1869 and states that it forms part
of his collection, I would have expected to find the material in the IRSNB (see
p. 271). However since Van Zwaluwenburg made this designation after examining
both collections I have accepted his decision.

Agrypnus melancholicus (Candéze) comb. n.

Lacon melancholicus Candéze, 1890 : CXLVIII.
Adeloceva melancholicus (Candéze) Fleutiaux, 1926 : 98.

LECTOTYPE (present designation). Inp1a: 9, Nagpore; melancholicus Cand.
| Nagpore, P.C. [Candéze, yellow border]; Lacon melancholicus Cd., det. E. Candéze
. | [IRSNB curatorial label]; Collection E. Candéze (IRSNB, Brussels).

| Paralectotype: 2, Nagpore; Lacon melancholicus Cd. det. E. Candéze [I[RSNB
| Curatorial label]; Collection E. Candéze (IRSNB, Brussels).

182 Cc. M. F. von HAYEK

Candéze based his description on four specimens. I have been unable to trace
the remaining two paralectotypes.

Candéze (1892 : 484) recorded this species as rare in Konbir, but found in large
numbers at Barwai. The IRSNB, Brussels collection contains a single female
labelled Barwai, Chota Nagpore, 1892 and 10 females from Nagpore with IRSNB
curatorial labels Lacon melancholicus Cd. det. E. Candéze. These specimens
are not conspecific with the lectotype. They are all Agrypnus truncatus (Candéze),
See p. 223).

Agrypnus meridionalis (Fleutiaux) comb. n.
Adelocera (Archontas) meridionalis Fleutiaux, 19346 : 58.
LECTOTYPE (present designation). MapaAGAscar: 9, Madagascar, Reg. Sud
de l’Isle; Ihosy, 2.33, A. Seyrig; meridionalis Fleut. type [Fleut.] (MNHN, Paris).

Paralectotypes. II ex., with the same locality labels as the lectotype (MNHN,
Paris). 1 9, Madagascar; Ihosy; Adelocera (Lacon) meridionalis Fleut. [Fleut.]
(BMNH).

Agrypnus minimus (Candéze) comb. n.

Lacon minimus Candéze, 1895 : 57.
Adelocera minimus (Candéze) Fleutiaux, 1926 : 96.

Holotype. Mapacascar: g, Madagascar, Diego Suarez, Ch. Alluaud; Museum
Paris, Madagascar, Coll. Ch. Alluaud 1904; L. minimus n.sp. Cand. det. [Cand.]
(MNHN, Paris).

Agrypnus minor (Candéze) comb. n.

Lacon minor Candéze, 1857 : 126.
Adeloceva minoy (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MApaAGAScAR: ¢, Madagascar; Janson
coll. ex Candéze 1903.130; Lacon minor Cdz. Mad. [Cand.] (BMNH).

Paralectotypes. 2 3, 1 9, Dej. Madagascar; Janson coll. 1903.130; Lacon minor
Cand. e coll. Dejean [Janson]. One specimen bears the word ‘Type’ on Janson’s
determination label (BMNH).

Candéze comments that there is material belonging to this species standing
as pumilo KI]. (in litt.) in the ZMHU, Berlin, and as gibbicollis (in litt.) in the
Chevrolat collection (MNHN, Paris). I have not examined these specimens.

Agrypnus minutus (Schwarz) comb. n.

Lobotarsus minutus Schwarz, 18984 : 133.

The description is based on an unrecorded number of specimens collected in
KAMERUN by Conrad and submitted by Kraatz.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 183

Type-material: ?DEI, Eberswalde.

The generic attribution is based on the following specimen: 2 SIERRA LEONE,
Rhobomp, determined by Fleutiaux (MNHN, Paris).

Agrypnus mixtus (Candéze) comb. n.

Lacon mixtus Candéze, 1888 : 669.
Adelocera mixtus (Candéze) Fleutiaux, 1926 : 96.

The description is based ona single specimen collected at Kawkareet, Tenasserim
{[BuRMA] in May by Fea.

Holotype. MCSN, Genoa (see p. 275).

The generic attribution is based on the description.

Agrypnus miyakei Ohira
Agrypnus miyakei Ohira, 1967) : 100.
Holotype. JAPAN: 3j, Amami-Oshima (Hatsuno), 30.iii.1964, Y. Miyake, Ohira
collection.

Paratypes. 2 9, ditto, 14.1V.1964, J. Nagao, Tokuno-Shima (Kamezu),
22.1V.1964, K. Arichi [sic]. This material is in the collection of the Entomological
laboratory, Ehime University and/or Nagao, Sat6, Shibata and Kishii collections.

I have not examined this material; the generic attribution is based on the
description.

Agrypnus miyakoensis (Ohira) comb. n.
Colaulon (Cryptolacon) miyakoensis Ohira, 1969a : 93.

Holotype. Japan: 9, Miyako-Jima, 26.VII.1965, H. Yamazaki, Ohira coll.
The generic attribution is based on the description.

Agrypnus miyamoti (Nakane & Kishii) comb. n.

Cryptolacon miyamoti Nakane & Kishii, 1955 : 2.
Colaulon (Cryptolacon) miyamoti (Nakane & Kishii) Kishii, 1964 : 23.

The description is based on a holotype, allotype and 50 paratypes from two
localities in the Tokara Islands [ Japan].

Type-material: Osaka Municipal Museum of Natural History, Entomological
Laboratory of the Kyushu University and in the authors own collections. Not
examined. The location of the individual specimens is not recorded but as the
authors record the data on each specimen there should be no difficulty in recognizing

them.

The generic attribution is based on the description.

184 Cos UE. “Won? (ELAS

Agrypnus mjobergi (Elston) comb. n.
Lacon mjobergi Elston, 1930 : 5.

The description is based on an unrecorded number of specimens from AUSTRALIA,
Queensland, Cedar Creek (Dr M7éberg).

Elston states that the type is in the NR, Stockholm but unless that museum
received only one specimen of the species this cannot be accepted as a valid
holotype designation. Neboiss (1952 : 10) records the presence of an unrecorded
number of paratypes in the AM, Sidney.

The generic attribution is based on Elston’s statement that the species resembles
L. marmoratus Candéze (=Agrypnus marmoratus (Candéze), see p. 181).

Agrypnus mocquerysi (Fleutiaux) comb. n.

Adeloceva mocquerysi Fleutiaux, 1932a : 50.

Holotype. Mapacascar: 2, Madagascar, Baie d’Antongil, A. Mocquerys: Type:
Adelocera mocquerysi Fleut., Type [Fleut.] (MNHN, Paris).

Agrypnus modestus (Candéze) comb. n.

Lacon modestus Candéze, 1857 : 118.
Adelocera modestus (Candéze) Fleutiaux, 1926 : 97.

LECTOTYPE (present designation). JAva: Dej. Java [Janson]; Janson coll.
1903 : 130; Lacon modestus Cdz. Cand. type (Agrypnus modestus Dej. coll.) e
coll. Dejean [Janson] (BMNH).

The description is based on material standing as Agrypnus modestus Dej. in
the de la Ferté Senéctére collection. The absence of Candéze’s determination

label and a label indicating that it came from the de la Ferté Sénectére collection
is probably due to Janson (see p. 276).

Agrypnus molestus (Candéze) comb. n.

Lacon molestus Candéze, 1857 : 120.
Adelocera molestus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Inp1a: g, Deyr. Dinapore, Ind. Or.
[Janson]; 289; Janson coll. 1903.130; L. molestus Cand. type [Cand.]; Lacon
molestus Cdze e coll. Deyrolle [Janson] (BMNH).

The published locality is ‘Dinapore, aux Indes-Orientales’ (see p. 271). There
is a town of this name in the State of Bihar. In this case ‘Indes-Orientales’ may

be a misprint for ‘Inde orientale’. There are no subsequent records of this species. |

—_—————____—_#_s_—___ hn |

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 185

Agrypnus molitor (Candéze) comb. n.

Lacon molitoy Candéze, 1875 : CXIX.
Adelocera molitoy (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Mindanao;
cote S—O [PHILIPPINES].
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on Candéze’s comment that this species resembles
setiger, judex, multiforis and fibrinus (q.v.).

Agrypnus mozambicana (Fleutiaux) comb. n.
Adelocera (Lacon auct.) mozambicana Fleutiaux, 19326 : 3.

Holotype. MozampBiguE: g, Museum Paris, Mozambique, Beira, P. Lesne,
1928; Decembre; Adelocera (Lacon) mozambicana Fleut. type [Fleut.] (MNHN,
Paris). This is an Agrypnus species, despite the fact that Fleutiaux compares it
with Adelocera aethiopicus (Candéze).

Agrypnus mucoreus (Candéze) comb. n.
Tilotaysus [sic] mucoreus Candéze, 1857 : 173.

LECTOTYPE (present designation). MApAGASCAR: ¢, 16084 [museum catalogue
number, refers to locality and collector on following label]; Madagascar, Goud.;
mucoreus Cand. Madag. Goud. [Gerstaecker] (NMHU, Berlin).

The absence of Candéze’ determination label is probably due to Gerstaecker
(see p. 275). The lectotype measures 15-5 mm compared with the published
length of 18 mm.

Paralectotypes: 1 g, 1 9, Madagascar Goud., No. 16084 (NHHU, Berlin). 1 J,
Madagascar; Janson Coll. 1903.130; Tilotarsus mucoreus Cdze. ex coll. de Mniszech
[Janson] (BMNH). The absence of Candéze’s determination label on the BMNH
specimen is probably due to Janson (see p. 276). The males measure 17 mm
and the female 16-5 mm.

Fleutiaux (1899 : 223) states that pulverews Candéze is an immature specimen
of this species but comparison of the lectotypes has shown that this is not the case.

Agrypnus multiforis (Candéze) comb. n.

Lacon multiforis Candéze, 1857 : 117.
Adelocera multiforis (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). E. PAKISTAN: 9, Sylhet [Janson]; Janson
coll. 1903.130; Lacon multiforis Cdz. Sihlet [sic, Cand.]; Lacon multiforis Cdze
Type (ex coll. Cand.) [Janson] (BMNH).

The published locality is ‘Indes-Orientales’ see p. 271.

186 cM. EF. von HAYEK

Agrypnus murinus (Linnaeus)

Elatey murinus Linnaeus, 1758 : 406, no. Ig. ;

Agrvypnus murinus (Linnaeus) Eschscholtz, 1829 : 32.

Lacon murinus (Linnaeus) Germar, 1840 : 264.

Adelocera mucorea LeConte, 1853 : 491. [Synonymized by Buysson, 1893 : 20.]
Lacon sordidus Candéze, 1857 : 114. [Synonymized by Fleutiaux, 1926 : 99.]
Agrypnus murinus (Linnaeus); Lane, 1953 : 88.

Elater murinus Linnaeus. LECTOTYPE (present designation). 9, 1g murinus
[Linnaeus] (Linnean Society collection, BMNH). The published locality is
‘EUROPE’.

The Linnean collection contains two additional specimens, both correctly
identified. The first, a male bears a printed number ‘28’ presumably referring to
Linnaeus, 1788 : 655 in which murinus is listed as number 28. The third specimen
bears a manuscript label ‘Angl. J.E.S.’ and was presumably added by J. E. Smith,
one-time owner of the Linnean collection.

Examination of the material standing as murinus Linnaeus in the BMNH,
IRSNB, Brussels and MNHN, Paris has shown that collectors and authors have
interpreted the species correctly. It is the only Agrypnus species occurring in
western Europe.

Adelocera mucorea LeConte. Holotype. 6, brick-red paper disk [Southern
States, see p. 277]; Lacon mucoreus Lec. [LeC., sic]; murinus (MCZ, Harvard).

The published locality is GzorciA. The contents of the abdomen have been
eaten by Anthrenus; the sex of the specimen has been determined from the
appearance of the last abdominal sternite (see p. 117). Why LeConte labelled
the specimen Lacon mucoreus when he described it as an Adelocera species is
unknown. As there is no detailed record of where the specimen was found, there
is no means of telling whether this specimen was imported by chance or whether
the brick-red paper disk was affixed to a European specimen in error. There
are no subsequent records of this species being found in the United States.

Lacon sordidus Candéze. LECTOTYPE (present designation). SENEGAL: g,
Collection Chevrolat; E. sordidus Cand. type [?Cand.]; sordidus Cand. type Mon.
[Fleut.]. The specimen stands over a Chevrolat collection label; Lacon sordidus
Cand. type Mon. I, p. 114 : 22, Senegal, Collection Chevrolat (MNHN, Paris).

The origin of this specimen is also unknown. There are no subsequent records
of this species from Senegal.

GERMAR’S INTERPRETATION OF Elater murinus LINNAEUS. The BMNH possesses
six specimens of this species from the Schaum collection. One bears a label
‘murinus omn.’ in a handwriting I believe to be that of Germar. All the specimens
are correctly determined.

Agrypnus murrayensis (Blackburn) comb. n.

Lacon murvayensis Blackburn, 18914 : 504.
Lacon murrayensis Blackburn; Neboiss, 1956 : Io.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 187

The description was based on an unrecorded number of specimens from
S. AUSTRALIA, near Murray Bridge.

The BMNH does not possess any specimens of this species. Neboiss (1956 : 10)
records that the type is in the SAM, Adelaide, but this cannot be accepted as a
valid holotype designation.

The generic attribution is based on the description.

Agrypnus muscerda (Candéze) comb. n.

Lacon muscerda Candéze, 1874 : 57.
Adelocera muscerda (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Cap de
Bonne—Espérance. (Coll. Cand.) [SoutH Arrica].

Type-material: ? IRSNB, Brussels.

The generic attribution is based on the following specimens, which may be
syntypes acquired by Janson. SouTH AFRICA: I ex., Janson coll. ex Candéze,
1903.130; Janson coll. 1903.130; L. muscerda Cdz. type [Cand.]. The abdomen
is missing. I 9, P.B.S. [Promontorium Bonae Spei]; Janson coll. 1903.130;
muscerda Cdz. [Cand.]; Lacon muscerda Cand. [Gahan]. 1 9, P.B.S.; Janson
coll. 1903.130; Lacon muscerda [Gahan] (BMNH).

The wings and metasternum are much reduced in length.

Agrypnus muscosus (Candéze) comb. n.

Lacon muscosus Candéze, 1893¢ : 170.
Adelocera muscosus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Kanara
and Belgaum in the Andrewes collection (see p. 270).

Syntypesexamined. INp1IA:1 J, Belgaum P.; Andrewes Bequest B.M. 1922.221;
Lacon muscosus Cand. [Andrewes]. 1 9, Belgaum S.; Andrewes Bequest B.M.
1922.221, 1 9, Canara; Andrewes Bequest B.M. 1922.221. 1 g, Kanara; Andrewes
Coll. B.M. 1933.109. I 4g, Kanara; co-type; B.M. 1946.93 [from Bishop coll.]
Lacon muscosus (BMNH).

The designation of a lectotype is postponed until any material extant in the
Candéze collection in the IRSNB, Brussels has been examined.

Agrypnus musculus (Candéze) comb. n.

Lacon musculus Candéze, 1857 : 141.
Colaulon (Cryptolacon) musculus (Candéze) Chujo, 1959 : 4.

The description is based upon an unrecorded number of specimens from CHINA
submitted by Dohrn.

188 C. M. F. von HAYEK

Type-material: 2I[PZAN, Warsaw (Dohrn coll.).

The generic attribution is based on the following specimen: J, China; Janson
coll. ex Candéze, 1903.130; Lacon musculus Cdz. China [Cand.]; Lacon musculus
Cand., ex. coll. Candéze [Gahan] (BMNH). This specimen may be part of the
syntype-series retained by Candéze.

Agrypnus mustellinus (Germar) comb. n.

Lacon mustellinus Germar, 1840 : 264.
Lacon birmanicus Candéze, 1874: 71. Syn.n.
Adelocera mustellinus (Candéze) Fleutiaux, 1926 : 96.

Candéze (1857 : 110) records that he received the ‘exemplaire typique de Germar’
of this species from Schaum. The BMNH manuscript catalogue of the Schaum
collection does not list this species. However the MNHU, Berlin contains a single
specimen from the Schaum collection (see below). Why this specimen is in Berlin
while the type-material of species described at the same time (tubidus Getmar
and tevrenus Germar) are now in the BMNH collection is unknown. There are
no specimens determined as mustellinus by Germar in the DEI, Eberswalde.

Lacon mustellinus Germar. LECTOTYPE (present designation). ¢, 43275
{Museum catalogue number = Siam, ex Schaum coll.]; L. mustellinus Germ.
[Candéze] (NMHU, Berlin). The specimen stands beside a yellow label in
Gerstaecker’s handwriting: mustellinus Germ. sec ejus type, Tenasserim Helf.
The published locality is Stam. The locality Tenasserim refers to a second specimen
of the same species bearing the number 16113. The absence of Germar’s determina-
tion label is probably due to Gerstaecker.

The lectotype agrees well with the description except that the antennae are
I1-segmented and not 12-segmented as stated by Germar. The last antennal
segment is a contricted at the apical third and this may have misled Germar,
whose eyesight was not very good (see p. 275), into believing that the antennae
were 12-segmented.

Lacon birmanicus Candéze. LECTOTYPE (present designation). BuRMA: g,
India, Rangoon; Janson coll. 1903.130; birmanicus Cdz. [Cand.]; Lacon birmanicus
Cand. type [Arrow] (BMNH).

Paralectotypes. 2 4, India, Rangoon; Janson coll. 1903.130 (BMNH).

Agrypnus muticus (Herbst) comb. n.

Elater muticus Herbst, 1806 : 93; pl. 166, fig. Io.
Lacon muticus (Herbst) Germar, 1840 : 262.
Adelocera muticus (Herbst) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). 3, Nr. 16126 (museum catalogue number =
data on next label): India or. Mus. Hbst.; (NMHU, Berlin).

Paralectotypes. I 4, data as lectotype. 1 9, Hbst.; 16126; India or. Mus.
Hbst. Nr. 16126. 19, Siam, Kaden; India or. Mus. Hbst. Nr. 16126. 1 9, Bintang

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 189

Rottgor; India or. Mus. Hbst. Nr. 16126. 2 9, India or. Mus. Hbst. Nr. 16126
(MNHU, Berlin).

The specimens stand beside a yellow label in Gerstaecker’s handwriting: muticus
Cand.,+ Elater muticus Hbst.+ Ind. or. The absence of Herbst’s determination
label is probably due to Gerstaecker (see p. 275). The published locality is ‘Ost
Indien’.

GERMAR’S INTERPRETATION OF Elater muticus HERBST. Germar refers to
Herbst, 1806 : 93; tab. 166, fig. 10 but as he records that the specimens before
him probably came from Arabia (corrected, loc. cit. p. 440, to Siam) it can be
assumed that he had before him material determined from the description. For
the history of the Germar collection see p. 275. Neither the DEI, Eberswalde,
or NMHU, Berlin contain this material. Candéze (1857: 154) states that he
received Germar’s specimens from Schaum. The BMNH manuscript catalogue
of the Schaum collection lists two specimens of muticus Helf. [sic] from Arabia.
The BMNH collection contains two female specimens without localities from the
Schaum collection. One bears a label: L. muticus Hbst. collect. Germ. in Candéze’s
handwriting. I believe that these specimens, which are correctly identified, are
those on which Germar based his interpretation of muticus Herbst.

Agrypnus myamoti (Nakane & Kishii) comb. n.

Cryptolacon myamoti Nakane & Kishii, 1955 : 1; pl. 1, fig. 1.
Colaulon (Cryptolacon) myamoti (Nakane & Kishii) Kishii, 1961 : 25.

Holotype. JAPAN: 3g, Takarajima 29.V.1953, T. Nakane leg. Allotype: 9,
Takarajima, 31.V.1953, IT. Nakane leg.

Paratypes: 41 ex, Takarajima, 26—30.V.1953. 9 ex, Nakanoshima, 4—13.VI.1953,
S. Miyamoto, T. Nakane & S. Uéno leg. The type-material is in the Osaka
Municipal Museum of Natural History, the Entomological Laboratory of the
Kyushu University and in the Nakane and Kishii collections.

Agrypnus mysticus (Candéze) comb. n.

Lacon mysticus Candéze, 1857 : 104.
Adelocera mysticus (Candéze) Fleutiaux, 1926 : 96.

Holotype. Mapacascar: 9, Laf. Madagascar; Madagascar; Janson coll.
1903.130; L. mysticus type [Cand.]; Lacon mysticus Cdze. Cand. Type e coll. de
Laferté [Janson]; 38 bis (BMNH).

Agrypnus nagaoi (Ohira) comb. n., sp. rev.
Colaulon (Sagojyo) nagaot Ohira, 1966 : 7.

The description is ‘based on a holotype and nine paratypes from the Ryukyu
IsLaANbs, Japan. The location of the material is not recorded. It is probably

190 CL Ma ES Vion; HAYEK

in the author’s collection or one of the collections recorded by Ohira (1966: 7,
footnote).

The generic attribution is based on the description.

Ohira (1971c : 66) synonymizes this species with lupinosus (Candéze) (p. 178)
but as the figure of the aedeagus of nagaoz (Ohira, 1971c : 65, fig. 2) shows it to
be very different from that of Jwpinosus the synonymy cannot be accepted.

Agrypnus ngokoensis comb. n.

Lobotarsus crenatus Schwarz, 1902a : 313.
Agrypnus crenatus (Schwarz) comb. n. [Secondary junior homonym of Agrypnus crenatus

(Klug, 1833).]

LECTOTYPE (present designation). CAMEROUN: 9, Cameroun, Vallee de la
N’Goko, Janvier 1900, Jobit; Lobotarsus crenatus Schw. [Schwarz] (MNHN,
Paris).

The description is based on an unrecorded number of specimens received from
Fleutiaux. There are no specimens from the Schwarz collection in the DEI,
Eberswalde.

Agrypnus niger (Schwarz) comb. n.

Lacon niger Schwarz, 1905 : 258.
Adelocera niger (Schwarz) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). InpIA: 9, Darjeeling: Coll. Schwarz;
Typus (DEI, Eberswalde).

Paralectotypes: 1 9, Darjeeling; coll. Schwarz; Typus. 2 2, same labels, and
also niger Schw. [Schwarz] (DEI, Eberswalde).

Agrypnus nigritus (Candéze) comb. n.

Lacon nigrita Candéze, 1857 : 137; pl. 2, fig. 7.
Adelocera nigrita (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). SouTH Arrica: Deyr. P.B.S. [Promonto-
rium Bonae Spei]; 308; Janson coll. 1903.130; L. nigrita Cdz. type [Cand.]; Lacon
nigrita Cdze. type e coll. Deyrolle [Janson] (BMNH).

The metasternum and wings are much reduced in length.

Agrypnus nivalis (Fleutiaux) comb. n.
Adelocera (Archontas) nivalis Fleutiaux, 1934) : 54.

LECTOTYPE (present designation). MADAGASCAR: 4, Madagascar, Perinet,
Forét cote est; nivalis Fleut. type [Fleut.] (MNHN, Paris).
Paralectotypes. xz 9, 2 ex., same locality labels as lectotype but without

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 191

determination labels 1 ex., Museum Paris, Madagascar, Mahatsinjo, Coll. J.
Chatanay 1914; nivalis Fleut. [Fleut.] (MNHN, Paris).

Agrypnus nodicollis (Candéze) comb. n.

Lacon nodicollis Candéze, 1857 : Iot.
Adelocera nodicollis (Candéze) Fleutiaux, 1926 : 96.

Holotype. Mapacascar: Madagascar; Janson coll. 1903.130; Lacon nodicollis
Cdze, type (e coll. de Mniszech) [Janson] (BMNH). The absence of Candeze’s
determination label is probably due to Janson (see p. 276).

The female from Madagascar, Diego Suarez, Alluaud, 1893 (Candéze, 1895 : 54,
locality) determined by Candéze in the IRSNB, Brussels is not conspecific with
the holotype. It belongs to a species unknown to me.

Agrypnus nodieri (Fleutiaux) comb. n.

Adelocera (Archontas) nodieri Fleutiaux, 1934) : 49.

: LECTOTYPE (present designation). MaApaGaAscar: 9, Diego Suarez, Madag.;
| Coll. Nodier; Adelocera nodieri Fleut., type [Fleut.] (MNHN, Paris).

Paralectotype. ¢, Museum Paris, Madagascar, Reg. de Sakarami, Maurice
de Rothschild, 1905; nodieri Fleut. [Fleut.] (MNHN, Paris).

Agrypnus nodifer (Klug) comb. n.

Elatey (Agrypnus) nodifer Klug, 1833 : 62; pl. 3, fig. 2 (1834 : 152).
Lacon nodifey (Klug) Germar, 1840 : 264.
Adelocera nodifer (Klug) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MADAGASCAR: @, 16092 [museum catalogue
_ number = data on the following label]; Madagascar Goud. (NMHU, Berlin). The
specimen stands beside a blue Gerstaecker label: nodifer Cand. Elater nodifer
Klug, Madag. Goudot. The absence of Klug’s determination label is probably
| due to Gerstaecker (see p. 275).

GERMAR’S INTERPRETATION OF Elater nodifer KiuGc. Germar does not give
a description of nodifer Klug. As neither the DEI, Eberswalde, NMHU, Berlin
nor BMNH collections contain any material bearing Germar’s determination label,
I have assumed that Germar based his interpretation of the species on Klug’s
| description or specimens.

Agrypnus nubilus (Candéze) comb. n.

| Tilotarsus [sic] nubilus Candéze, 1857 : 175.
| Lobotarsus nubilus (Candéze) Schwarz, 1906 : 30.

LECTOTYPE (present designation). GaxBon: 9, Gaboon; Janson coll. 1903.130;

192 C. M. F. von HAYEK

Tilotarsus nubilus Cdz. Gaboon Deyr. [Cand.]; Tylotarsus nubilus Cdze., ex coll.
Candéze [ Janson] (BMNH).

I have not found any specimens from Guinea and Benguela, the other two
localities recorded by Candéze.

Agrypnus oberndorferi (Schwarz) comb. n.

Lacon oberndorfert Schwarz, 19020 : 197.
Adelocera oberndorfevi (Schwarz) Fleutiaux, 1926 : 96.

The description is based on a holotype from Mapras, Dindigul submitted by
Obernderfer. This specimen cannot be found in the Schwarz collection in the
DEI, Eberswalde.

The generic attribution is based on the following specimen; BuRMA: 9, Rangoon;
coll. Schwarz; oberndorferi Schw. [Schwarz] (DEI, Eberswalde). The specimen
is 7mm long compared with the published length of 9 mm.

Agrypnus obscurus (Fleutiaux) comb. n.
Adeloceva (Archontas) obscura Fleutiaux, 1934) : 51.
LECTOTYPE (present designation). MADAGASCAR: 9, Madagascar, Lamberton,
obscura Fleut., type [Fleut.] (MNHN, Paris).
The published locality is Madagascar (Lamberton).

Agrypnus octavus (Candéze) comb. n.

Lacon octavus Candéze, 1874 : 95.
Adeloceva (Compsolacon) octava (Candéze) Van Zwaluwenburg, 1959 : 354.

Lectotype (designated by Van Zwaluwenburg, 1959: 354, see p. Q).
AUSTRALIA: 2, N. H. Holl. C. Cdze.; Janson coll. ex Candéze, 1903.130; Lacon
octavus Cdz. n.sp. N. Austr. Th. [Cand.]; Lacon octavus Cand. type, Queensland
[Gahan] (BMNH).

Candéze comments that in shape this species resembles Myrmodes [akidiformis
Candéze]. In both species the metathorax is much reduced in length and the
wings no more than minute flaps. The main difference is that in the Myrmodes
species the humera are less strongly angled.

Agrypnus orientalis Hope
Agrypnus orientalis Hope, 1843 : 63.

Both Candéze (1891) and the Schenkling Catalogue (1925) record the date of
publication of orientalis as 1845. However a shorter form of Hope’s 1845 paper
appeared (see Wheeler, 1912) in the Proceedings of the Entomological Society

°

|
|

;
,

-
,
'
7

;

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 193

for March 7th, 1842, which were published in the Amunals and Magazine of Natural
History in January, 1843.

LECTOTYPE (present designation). ?WeEstT Mataisia: 9, Penang/Dr Cantor;
orientalis Hope (BMNH). The specimen lacks a registration label but the locality
and identification labels are identical to those found on specimens in the BMNH,
known to come from the East-India Company’s museum. The locality is probably
erroneous (see notes on the Cantor collection, p. 272).

Candéze (1891 : 22) tentatively places this species in synonymy with bipapulatus
Candéze, 1865. Confirmation of the synonymy is postponed until more material
becomes available and the species can be studied in greater detail.

Agrypnus ornatellus (Candéze) comb. n.

Lacon ornatellus Candéze, 1889 : 72 (6).
Adelocera ornatellus (Candéze) Fleutiaux, 1926 : 99.

The description is based on an unrecorded number of specimens from MADAGASCAR.
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: 9, Madagascar
Goudot, 1834; ornatellus Cand. [Cand.] (MNHN, Paris).

Agrypnus ornatus (Candéze)

Lacon ornatus Candéze, 1857 : 102.
Agrypnus ornatus (Candéze) Ohira, 1971 : 226.

The description is based on an unrecorded number of specimens from JAVA in
the Mniszech collection.

Type-material: ?IRSNB, Brussels (see p. 273).

The generic attribution is based on the following specimen: JAvA; Lacon ornatus
Cdz., Java [Cand.]; Lacon ornatus Cand. ex coll. Cand. [Gahan] (BMNH).

Agrypnus oshimanus Ohira
Agrypnus (Agrypnus) oshimanus Ohira, 1969a : 91; fig. 1, pl. 1, fig. 13.

Holotype. JAPAN: gj, Amami-Oshima (Hatsuno), 3.1V.1964 (T. Akashi) (Ohira
collection).

Paratype. 9, same data as the holotype (Ohira collection).

Agrypnus paenulatus (Boheman) comb. n.

Lacon paenulatus Boheman, 1851 : 415.

| Adeloceva paenulatus (Boheman) Fleutiaux, 1926 : 96.

Archontoides pretoriensis Cobos, 1966 : 651. Syn. n.
N

194 Gi Ms Es VON VEAMIEKK

Lacon paenulatus Boheman. LECTOTYPE (present designation). SouTH
AFRICA: 4, Caffraria; J. Wahlb. (NR, Stockholm).

Paralectotypes. I ex (? g, abdomen missing), I 9, same labels as lectotype.
I ex. (? 9, abdomen missing). Caffraria; J. Wahlb.; paenulatus Boh. [Boh.]
(NR, Stockholm).

The published locality is ‘Habitat in terra Natalensi’. The localities of
Boheman’s material are discussed on p. 280. The male was selected as the
lectotype because the specimen bearing Boheman’s determination label lacks
the abdomen.

Archontoides pretoriensis. Cobos. Holotype. SouTH AFRIcA: Q,. Fountains,
Pretoria, Africa det. Sur (Al. Capner coll. 3.1.1951 leg); Archontoides pretoriensis;
Archontoides pretoriensis mihi [Cobos] (Cobos collection).

Agrypnus paleatus (Champion) comb. n.

Lacon paleatus Champion, 1894 : 267.
Pyrganus paleatus (Champion) Golbach, 1968 : 198.

Syntypes examined. PANAMA: 2 9, 49, V. de Chiriqui, 25-4000 ft. Champion;
BCA Coll. IlI(x) Lacon paleatus Ch. [Champion] (BMNH). Two males are
marked ‘9’ by Champion (see p. 117) and one female bears a label: ‘sp. figured’.
Lectotype designation is postponed until the specimens have been studied in

greater detail.
It is of interest to note that in all the syntypes the margins of the prothorax
are crenate to a greater or lesser degree (cf. Golbach’s generic diagnosis of Pyrganus).

Agrypnus palliatus (Candéze) comb. n.

Lacon palliatus Candéze, 18930 : 7.
Adeloceva pailiatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from MADAGASCAR,
Andrangoloaca.

Type-material: ?>IRSNB, Brussels.

The generic attribution based on Candéze’s comment ‘Jolie espéce du groupe
du turbidus’ (=Agrypnus tubidus (Germar), see p. 224).

Agrypnus palliditarsis (Candéze) comb. n.

Lacon palliditarsis Candéze, 1857 : 120; pl. 1, fig. 18.
Adeloceva palliditarsis (Candéze) Fleutiaux, 1926 : 99.

LECTOTYPE (present designation). SoutH Arrica: 9, Type; 106; L. palli-
ditarsus [Cand.]; Adelocera palliditarsis Chevr. Schaum [unknown mss., last few
words illegible]; Collection Chevrolat; n.sp. Germ. palliditarsus Cand. type Mon.
[Fleut.] (MNHN, Paris). The specimen stands beside a Chevrolat collection label: |
Lacon palliditarsis Cand., type Mon. 1 : 120, 33 Cap. bon. sp. Chevrolat.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 195

Agrypnus pallidus (Candéze) comb. n.

Lacon pallidus Candéze, 1887 : 190.
Adelocera pallidus (Candéze) Fleutiaux, 1926: 96.

Holotype. Sumatra: 2, Dr B. Hagen, Tandjong Morawa, Serdang (N. O.
Sumatra) L. pallidus Cdz. n.sp. [Cand.] (RNH, Leyden).

Agrypnus palpalis (Candéze) comb. n.

Lacon palpalis Candéze, 1882 : 13.
Adelocera (Compsolacon) palpalis (Candéze); Van Zwaluwenburg, 1959 : 354.

Lectotype (designated by Van Zwaluwenburg, 1959 : 354). AUSTRALIA: sex
undetermined, Northern Australia, from the Monchicourt collection (IRSNB,
Brussels). Not examined.

Van Zwaluwenburg does not record whether the specimen bears a locality or
determination label.

The published locality is ‘Australia Septentrionale: Cleveland Bay’ [near
Townsville, Queensland]. The fact that Candéze records that he found the
material in the ‘ancienne collection Monchicourt’ suggests that the material was
in his own collection. Candéze acquired the Monchicourt collection via E. Deyrolle
(Horn & Kahle, 1936: 181). Whether the two additional specimens recorded
by Van Zwaluwenburg formed part of the original syntype series is unknown.

Agrypnus pantherinus (Fleutiaux) comb. n.

Centrostethus pantherinus Fleutiaux, 19346 : 60.

LECTOTYPE (present designation). MaApaGascar: 9, Madagascar, Perinet,
Forét cote est.; pantherinus Fleut. type [Fleut.] (MNHN, Paris).

Paralectotype. I ex., same locality as the lectotype, without determination
labels (MNHN, Paris).

I have not seen specimens from the other four recorded localities, Fort Tanala
(Ch. Alluaud), Ifandana (coll. Ch. Alluaud). Mahatsinjo, prés Beforona (coll.
J. Chatanay), Province de Mananjary (Goissaud). This material should also be
in the MNHN, Paris.

Agrypnus parallelicollis (Candéze) comb. n.

Lacon parallelicollis Candéze, 1857 : 118.
Adelocera parallelicollis (Candéze) Fleutiaux, 1926 : 96.

Holotype. JAvA: 9, Java, L. parallelicollis Cand. [Cand.]; Collection Chevrolat;
parallelicollis Cand. type [Fleut.]. The specimen stands beside a yellow Chevrolat

_ label: Lacon parallelicollis Cand. type Mon. 1, p. 118,29. Java, Collection Chevrolat

(MNHN, Paris).

196 C. M. F. von HAYEK

Agrypnus parallelus (Candéze) comb. n.

Lacon parallelus Candéze, 1874 : 90.
Lacon parallelus (Candéze) ; Neboiss, 1956 : 11.

The description is based on an unrecorded number of specimens from Queensland.
Neboiss (loc. cit) states that the type is in the BMNH, but the date of publication
(see p. 271) suggests that the type-material should be in the IRSNB, Brussels.
Selection of a lectotype is postponed until any material in that collection has
been examined.

Generic attribution based on the following specimen, possibly a syntype;
AUSTRALIA: g, N. Holl. C. Cdze.; Janson coll. ex Candéze, 1903.130; Lacon
parallelus Cdz. n.sp. N. Austr. Th. [Cand.] Lacon parallelus Cand. type [Gahan]
(BMNH).

Agrypnus parcus (Schwarz) comb. n.
Tilotarsus [sic] parcus Schwarz, 1903a : 338.

The description is based on an unrecorded number of specimens from MADAGASCAR.
Type-material: ?>DEI, Eberswalde.
The generic attribution is based on the description.

Agrypnus parviceps (Gyllenhal) comb. n.

Elater parviceps Gyllenhal in Schoenherr, 1817 : 133.
Lacon parviceps (Schoenherr) Germar, 1840 : 263.

LECTOTYPE (present designation). SIERRA LEONE: g, S. Leone, Afzelius
(NR, Stockholm). The specimen lacks a determination label but as far as is
known the specimen has always stood and been regarded as the type of parviceps
Gyllenhal. The species is not listed in Schenkling (1925).

GERMAR’S INTERPRETATION OF Elater parviceps GYLLENHAL. Germar comments
that Elater parviceps Schénh. [sic] appears to be related to Lacon terrenus.
Whether he based his interpretation on the description or on determined specimens
is unknown, but examination of the material in the NR, Stockholm has shown
that he was justified in his belief. A. parviceps (Gyllenhal) differs from terrenus
Germar (=crenatus Klug) in that the fourth tarsal segments are lobed.

Agrypnus parvus (Fleutiaux) comb. n.

Centrostethus paruvus Fleutiaux, 19346 : 61.

LECTOTYPE (present designation). MApAGAscAR: 4, Madagascar, Perinet,
Forét cote est; parvus Fleut. type [Fleut.] (MNHN, Paris).

Paratypes. 4 ex, same locality as lectotype, without determination labels
(MNHN, Paris).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 197

Agrypnus pauliani (Girard) comb. n.
Archontas pauliani Girard, 1970 : 28.

Holotype. MApAGAscAR: 3g, massif du Tsaratanana, Matsabory en dessous
de l?Andohanisambirano, 1,900m [2,050 m], début xii.1964 (P. Soga) (MNHN,
Paris).

Paratype. 1 9, same locality as holotype (Girard collection, Laboratoire de
Zoologie, Ecole normale supérieure, Paris).

Both specimens bear Girard’s determination label.

Agrypnus pauper (Candéze) comb. n.

Lacon pauper Candéze, 1878a : 100.
Adelocera pauper (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Teibodas
JAVA, collected by Beccari.

Type-material: MCSN, Genoa and ?IRSNB, Brussels.

The generic attribution is based on the description.

Agrypnus perplexus (Elston) comb. n.

Lacon perplexus Elston, 1924 : 202.
Lacon perplexus Elston; Neboiss, 1956 : II.

The description is based on an unrecorded number of specimens from N.W.
AUSTRALIA: Forest River (W. Crawshaw), Hammersley Range (W. D. Dodd),
Wyndham (W. Crawshaw), Northern Territory: Port Darwin.

Syntype-material (see p. 274): AM, Sidney, SAM, Adelaide, NM, Victoria.

The generic attribution is based on syntype 2, Wyndham (W. Crawshaw) (SAM,
Adelaide).

Agrypnus pictilis (Schwarz) comb. n.

Lacon pictilis Schwarz, 1899 : 76.
Adelocera pictilis (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from D. Ost.
Africa [TANZANIA], collected by Bennigsen.
Type-material: >DEI, Eberswalde.

The generic attribution is based on the following specimen; KEnya: 2, Malanga,
Br. O. Afr.; pictilis Schw. comparé au type [Fleut.] (MNHN, Paris).

Agrypnus pictipennis (Candéze) comb. n.

Lacon pictipennis Candéze, 1857 : 150.
Lacon pictipennis Candéze; Neboiss, 1956 : II.

198 C. M. F. von HAYEK

The description is based on an unrecorded number of specimens from Van
Diemansland [Tasmania]. Candéze adds that he received the species from la
Ferté Sénectére, determined as Agrypnus pictipennis Gory. The BMNH does
not possess any specimens from Tasmania determined by Candéze. As Candéze
(1874 : 93) records the locality as ‘Nouvelle Galles du sud et Victoria’ and as
‘Austr. merid.’ in the catalogue (1891 : 25) it is possible that the originally published
locality is erroneous.

LECTOTYPE (present designation). AUSTRALIA: 9, Laf. Tasmania [Janson];
pictipennis Gory, Nov. Holl. [? Laferté Sénectére]; Lacon pictipennis Cdze Cand.
Type e coll. de Laferté [Janson]; Janson coll. 1903.130 (BMNH). The locality
label ‘Tasmania’ was obviously added by Janson following Candéze’s original

publication. The absence of Candéze’s determination label is almost certainly
due to Janson.

Agrypnus pictiventris (Candéze) comb. n.
Lacon pictiventris Candéze, 1857 : 127; pl. 2, fig. 8.
Adelocera pictiventris (Candéze) Fleutiaux, 1926 : 96.

Holotype. Mapacascar: ¢, Laf. Madagascar; pictiventris [Cand.]; Coquerel;

Lacon pictiventris Cdze, Cand. Type e coll. de Laferte [Janson]; Janson coll.
1903.130 (BMNH).

Agrypnus piger (Candéze) comb. n.
Lacon pigey Candéze, 1889 : 73 (7).
Adelocera pigery (Candéze) Fleutiaux, 1926 : 96.
The description is based on an unrecorded number [‘plusieurs’] of specimens
collected at Tetara, BENGAL by Renard.
Type-material: ?>[RSNB, Brussels.

The generic attribution is based on the following specimen, possibly a syntype:
9, Tetara; L. piger nov. spec. fasicule IV, p. 7 [Cand.] (MNHN, Paris).

Agrypnus pinguis (Candéze) comb. n.
Lacon pinguis Candéze, 18930) : 9.
Lacon pinguis Candéze; Neboiss, 1956 : II.
The description is based on an unrecorded number of specimens from AUSTRALIA,
Cooktown [Queensland].
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on the following specimen; AUSTRALIA: 9,
Austr.; pinguis Cand. det. Cand. teste Fleut. [Fleut.] (MNHN, Paris).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 199

Agrypnus pipitzi (Candéze) comb. n.
Lacon pipitzi Candéze, 1889 : 72 (6).
Adelocera pipitzi (Candéze) Fleutiaux, 1926 : 96.
The description is based on an unrecorded (‘plusieurs’) number of specimens
received from Pipitz.
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: MADAGASCAR: 3,
Madagascar: pipitzi Cand. [Fleut.] (MNHN, Paris).

Agrypnus pistorius (Candéze) comb. n.

Lacon pistorius Candéze, 1893¢ : 171.
Adelocera pistorius (Candéze) Fleutiaux, 1926 : 96.

Syntypes examined. INpIA: 2 J, 2 9, Kanara; cotype (BMNH curatorial label].
One specimen bears Andrewes’ determination label, and another a determination
label in an unknown hand (BMNH). The description was based on six specimens.
The selection of a lectotype is postponed until a search has been made for the
remaining two specimens.

Agrypnus plagiatus (Candéze) comb. n.

Lacon plagiatus Candéze, 1878b : LIV (8).
Lacon plagiatus Candéze; Neboiss, 1956 : 11.

The description is based on an unrecorded number of specimens from Australia,
Port Denison [near Bowen, QUEENSLAND].
Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: 2, Lacon plagiatus
Cand. Cooktown [?Cand.] (MNHN, Paris).

Agrypnus planatus (Candéze) comb. n.

Lacon planatus Candéze, 1895) : 55.
_ Adelocera planatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Mapa-
GASCAR, Imerina (Sikora).

Type-material: ?>IRSNB, Brussels.

The generic attribution is based on the following specimen: MADAGASCAR: g,
Madag., Collection Fleutiaux; L. planatus Cand. n.sp. El. n. vi [?Cand.], L. planatus
Cand. Ann. Belg. 95 : 55. Cand. det. [Fleut.] (MNHN, Paris).

The description did not appear in Elaterides Nouveaux 6, 1806.

200 Gi Me EB: von HAY Ek

Agrypnus pleureticus (Candéze) comb. n.

Lacon pleureticus Candéze, 1874 : 93.
Lacon pleureticus Candéze; Neboiss, 1956 : II.

The description is based on an unrecorded number of specimens from Queensland,
Rockhampton and Brisbane in the Candéze and Janson collections.

Syntypes examined. AUSTRALIA: I 3g, N. Holl./C. Cdze; Lacon pleureticus
Cdz. n.sp. N. Austr. Th. [Cand.]; Lacon pleureticus Cdz. ex coll. Cand. [Janson]
type [Gahan]. 1 3, same locality and collection, but without determination labels
(BMNH). The selection of a lectotype is postponed until any material from the
Candéze collection extant in the IRSNB, Brussels has been examined.

Agrypnus porcinus (Candéze) comb. n.

Lacon porcinus Candéze, 1857 : 143.
Adelocera porcinus (Candéze) Fleutiaux, 1926 : 96.

Holotype. Reunion: g, Laf. Insula Bourbon; Janson coll. 1903.130; Lacon
porcinus Cdze. Cand. type e coll. de Laferté [Janson]; porcinus Cdz. type [Cand.]
(BMNH).

Agrypnus porosus (Klug) comb. n.

Elater (Agrypnus) porosus Klug, 1835 : 65 (1834 : 153).
Adelocera porosus (Klug) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MADAGASCAR: g, 16099 [museum catalogue
number refers to data on following label]; Madag. Goud.; porosus Cand. Elater
porosus Klug Madag. Goud. [Gerstaecker] (NMHU, Berlin).

Paralectotype. 3, Madagasc. Goud. Nr. 16099 (NMHU, Berlin).

Agrypnus porrectus (Fleutiaux) comb. n.

A deloceva (Archontas) porvecta Fleutiaux, 19346 : 58.

LECTOTYPE (present designation). MADAGASCAR: 3, Fianarantsoa Mada-
gascar; porrecta Fleut. type [Fleut.] (MNHN, Paris).

Agrypnus porriginosus (Candéze) comb. n.

Lacon porriginosus Candéze, 1874 : 93.
Lacon porriginosus Candéze; Neboiss, 1956 : II.

LECTOTYPE (present designation). AUSTRALIA: 2, W. Australia; Nikol Bay;
Janson coll. 1903.130; L. porriginosus Cdz. [Cand.]; Lacon porriginosus Cand.
type, Nikol Bay [Gahan] (BMNH).

)
|
)

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 201

Agrypnus praelongus (Elston) comb. n.

Lacon praelongus Elston, 1927 : 355.
Lacon praelongus Elston; Neboiss, 1956 : II.

The description is based on an unrecorded number of specimens from Cunnamulla
[Queensland] (H. Hardcastle).
Syntype material (see p. 274). SAM, Adelaide, AM, Sydney.

The generic attribution is based on the following specimen: AUSTRALIA: Queens-
land, Dr K. K. Spence; determined by Carter, teste Fleutiaux [Fleut.| (MNHN,
Paris).

Agrypnus praetermissus (Candéze) comb. n.
Lacon praetermissus Candéze, 1874 : 76.

Adelocera praetermissus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Inp1A: 3, India [Janson]; Janson coll.
1903.130; Lacon praetermissus Cand. type India [Gahan]; sp. nov. ac distinctissima
nec serrula sec typ. et descript. L. praetermissus mihi [Janson]; Lacon serrula
Cdz. [Cand.]; Lacon serrula [crossed out] Cdze Cand. e coll. D. Parry [Janson]
(BMNH).

The description is based on an unrecorded number of specimens from ‘Indes
orientales (Coll. Janson)’. The absence of Candéze’s determination label and
the discrepancy between the published locality and the label are probably due
to Janson (see p. 276).

Agrypnus praetexta (Fleutiaux) comb. n.

Adelocera (Archontas) praetexta Fleutiaux, 1934) : 57.

LECTOTYPE (present designation). MapacAscar: g, Madagascar, Perinet,
Forét cote est.; Type; praetexta Fleut., type [Fleut.] (MNHN, Paris).

Agrypnus princeps (Fleutiaux) comb. n.

Lacon princeps Candéze, 1874 : 89.
Adelocera (Compsolacon) princeps (Candéze); Van Zwaluwenburg, 1959 : 354.

Lectotype (designated by Van Zwaluwenburg, 1959 : 354). 9, AUSTRALIA:
Cape York; Janson coll. 1903.130; L. princeps Cdz. type [Cand.]; Lacon princeps
Cand., type, Cape York [Gahan] (BMNH).

Agrypnus procellosus (Candéze) comb. n.

Lacon procellosus Candéze, 1895b : 54.

| Adelocera procellosus (Candéze) Fleutiaux, 1926 : 99.

202 Co Mi Ss Von ELA GE TG

The description is based on an unrecorded number of specimens from MApDA-
GASCAR, Imérina (Sikora).

Type-material: ?IRSNB, Brussels.

The generic attribution is based on the following specimen: 9, Madag.; L.
procellosus Cand. El.n.vi [?Cand.]; L. procellosus Cand. Ann. Belge 1895 p. 54
Cand. det. [Fleut.] (MNHN, Paris).

Agrypnus proxima (Fleutiaux) comb. n.

Adelocera (Archontas) proxima Fleutiaux, 1934) : 54.

LECTOTYPE (present designation). Mapacascar: 4, Diego Suarez; proxima
Fleut., type [Fleut.] (MNHN, Paris).

Paralectotype: I g, Diego Suarez; proxima Fleut. [Fleut.] (MNHN, Paris).

Agrypnus pulvereus (Candéze) comb. n.
Tilotarsus [sic] pulvereus Candéze, 1889 : 74 (8).
LECTOTYPE (present designation). MaApacascar: 9, Museum Paris, Mada-

gascar, Boivin 12.53; 12.53, Madagascar, M. Boivin; Tilotarsus pulvereus Cand.
sp.n. 89 [Cand.]; mucoreus [Fleut.] (MNHN, Paris).

Fleutiaux (1899 : 223) states that this is an immature mucoreus Candéze, but
comparison of the lectotypes has shown that this is not the case.

Agrypnus punctatissimus (Elston) comb. n.

Lacon punctatissimus Elston, 1927 : 362.
Lacon punctatissimus Elston; Neboiss, 1956 : 12.

The description is based on an unrecorded number of specimens from AUSTRALIA;
N. Territory, Groote Eylandt (N. B. Tyndale).
Syntypes (see p. 274). SAM, Adelaide, AM, Sydney.

The generic attribution based on the following syntype; AUSTRALIA: 9, Groote
Eylandt, N. Territory (N. B. Tyndale) (SAM, Adelaide).

Agrypnus punctipennis (Candéze) comb. n.
Lacon punctipennis Candéze, 1874 : 99.

Lacon punctipennis Candéze; Neboiss, 1956 : 12.

LECTOTYPE (present designation). AUSTRALIA: g, West Austral.; Janson
coll. 1903.130; punctipennis Cdz. type [Cand.]; Lacon punctipennis Cand. type
W. Australia [Janson] (BMNH).

Paralectotype. 1 9, West Australia/Nikol Bay; Janson coll. 1903.130; Lacon
punctipennis Cand. cotype [Gahan] (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 203

The description is based on an unrecorded number of specimens from ‘Australie
occidentale (Coll. Janson)’. The wings and metasternum are much reduced in
length.

Agrypnus pupillus (Candéze) comb. n.

Lacon pupillus Candéze, 1892b : 802.
Adelocera (Compsolacon) pupilla (Candéze); Van Zwaluwenburg, 1959 : 354.

Lectotype (designated by Van Zwaluwenburg, 1959 : 354). NEW GUINEA: 79,
Dilo, New Guinea (Loria) (MCSN, Genoa). Van Zwaluwenburg does not record
whether the specimen bears Candéze’s determination label.

Paralectotypes. Van Zwaluwenburg states that the lectotype is the first of
seven specimens, but does not record the data on the remaining six. Whether
they are paralectotypes can be determined only by inspection of the locality labels.
7 ex, N. Guinea, Dilo, Loria VI-VII.go0; Coll. R. Oberthur, 1952; 2 specimens
are labelled: Lacon pupillus n.sp. Cand. [?Cand.]. 5 ex, same locality, one
specimen with a similar determination label. 1 ex, N. Guinea Ighibirei, Loria
Vii.viii.go; L. pupillus Cand. n.sp. [?Cand.]; Cand. Ann. Mus. Civ. Gen. 1892 p. 80
[Fleut.] (MNHN, Paris). The specimens recorded from Hula, etc., by Candéze
have not been located.

Agrypnus quadricollis (Fairmaire) comb. n.

Lacon quadricollis Fairmaire, 1903 : 202.
Adelocera quadricollis (Fairmiare) Fleutiaux, 1926 : 99.

Syntypes examined. MADAGASCAR: I ex, Madag. Perrier; Museum Paris,
Madagascar, Perrier de la Bathie 1906; Lacon [Fleut) quadricollis Fairm. [Fairm]
Fleutiaux det., 4 ex, same locality and collection without determination labels. 1 ex,
ibidem; quadricollis Fairm, ex typ. [Fleut.] (MNHN, Paris).

The metasternum and wings are much reduced in length.

Agrypnus quadrinotatus (Schwarz) comb. n.

Lacon quadrinotatus Schwarz, 1908a : 73.

Lacon quadrinotulus Schenkling, 1925 : 32. [Unnecessary replacement name, see Schenkling,
O27 : 517.]

Adelocera quadrinotatus (Schwarz) Fleutiaux, 1926 : 99.

The description is based on an unrecorded number of specimens from Madagascar.
Type-material: ? DEI, Eberswalde.

The generic attribution is based on the following specimen: MADAGASCAR, 3,
Nossi bé, Loucoube, Ch. Alluaud, 1897; Lacon ase itns Schwarz, comparé au
type [Fleut] (MNHN, Paris).

204 GC. M. EF. von HAYEK

Agrypnus quedenfeldti nom. n.

Tylotarsus fuscus Quedenfeldt, 1886 : 25.
Agrypnus fuscus (Quedenfeldt) comb. n. [Junior secondary homonym of Agrypnus fuscus
(Fabricius 1801).]

LECTOTYPE (present designation). 4, Tylotarsus fuscus Qdf. Ogo. [Qued];
Ex Museum Quedenfeldt (MNHN, Paris).

The published localities are ANGOLA and Quango R. [Kwango or Cuango] Coll.
Mechow. The exact locality is unknown.

Agrypnus rectangularis (Say) comb. n.

Elater vectangularis Say, 1825 : 263.
Colaulon vectangularis (Say) Arnett, 1952 : 118.

The description is based on an unrecorded number of specimens ‘I found near
the Rocky Mountains during the journey of Major Long’s exploring party’. From
the account of this exploring party (James, 1823) the type-locality is probably
eastern Colorado, between 20°20’N. and 40°40’N. and 104°W. and 105°W. As Say
almost certainly retained the material he collected, it has presumably been lost
together with the rest of his collection (see p. 279). Miss P. Schuyler reports that
it cannot be found in the ANS, Philadelphia. Long’s material has not been located
(see p. 278).

The generic attribution is based on material standing over this name in the
BMNH and MNHN, Paris. The BMNH series includes specimens with wings of
normal length and others in which the wings are much reduced. Examination of
series of specimens may show that a reduction in wing length, unaccompanied by
a reduction in the length of the metathorax is an intraspecifically variable charac-
teristic. However, until a neotype is designated, the identity of vectangularis Say
remains unknown.

For Adelocera curtus Le Conte which Arnett (1952 : 118) and Golbach (1968 : 198)
regard as a synonym of rectangularis see p. 146.

Agrypnus recticollis (Elston) comb. n.

Lacon recticollis Elston, 1930 : 4.

The description is based on an unrecorded number of specimens from AUSTRALIA;
Queensland, Malanda (Dr E. Mjoberg). Elston states that the type is in the NR,
Stockholm, but unless that museum received only one specimen, this statement
cannot be accepted as a valid type-designation. Neboiss (1956: 12) records the
presence of paratypes in the AM, Sidney.

The generic attribution is based on the description and Elston’s statement that
the species resembles vandepolli Candéze (= Agrypnus vandepolli, see p. 227).

At first sight Lacon recticollis Elston appears to be a junior primary homonym of
Lacon recticollis Fleutiaux, 1927. However, the two species are not congeneric as

LL LL

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 205

Fleutiaux’s interpretation of Lacon is that of Castelnau, 1836 (and the present work»
type-species punctatus Herbst) while Elston, apparently unaware of Fleutiaux’s
(1926) correction of the Schenkling catalogue, continued to use Germar’s (1840)
interpretation of Lacon (type-species murinus Linnaeus = Agrypnus Eschscholtz
of the present work).

Agrypnus reductus (Candéze) comb. n.
Lacon rveductus Candéze, 1878b : LIV (8).
Adeloceva veductus (Candéze) Fleutiaux, 1926 : 96.
The description is based on an unrecorded number of specimens from SUMATRA.
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on Candéze’s observation that the species bears
a close resemblance to fibrinus Candéze (= Agrypnus, see p. 155).

Agrypnus reticulatus (Elston) comb n.
Lacon reticulatus Elston, 1930 : 6.
~
The description is based on an unrecorded number of specimens from N. W.
AUusTRALIA, Kimberley District, Nomkanbah (Dr FE. Mjoberg). Elston states that
the type is in the NR, Stockholm but unless that museum received only one specimen,

this cannot be accepted as a valid lectotype designation. Neboiss (1956: 12)
records the presence of paratype material in the AM, Sidney.

The generic attribution is based on the description and the fact that Elston
compares the species with Lacon variabilis Candéze (=Agrypnus, see p. 227).

Agrypnus rohanchaboti (Fleutiaux) comb. n.
Tylotarsus vohanchaboti Fleutiaux, 1922b : 45.
LECTOTYPE (present designation). ANGOLA: ¢g, Museum Paris, Angola

Benguela Capelongo—Dongo, Mission Rohan Chabot 1914; Tylotarsus rohanchaboti,
Fleut., type [Fleut.] (MNHN, Paris).

The second specimen mentioned in the description has not been located.

Agrypnus rubiginosus (Candéze) comb. n.

Lacon rubiginosus Candéze, 1882 : 11.
Lacon rubiginosus Candéze; Neboiss, 1956 : 12.

LECTOTYPE (present designation). AUSTRALIA: 2, Austr. sept.; L. rubiginosus
Cdz., type [Cand.]; ex Museo van Lansberge (MNHN, Paris).

The description is based on an unrecorded number of specimens in the Lansberge
collection from Australia septentrionale; Baie de Darwin.

206 C. M. F. von HAYEK

Agrypnus rufus (Blackburn) comb. n.
Pseudolacon rufus Blackburn, 1890 : 90.

LECTOTYPE (present designation). AUSTRALIA: 3, 3371 [probably refers to
SAM, Adelaide catalogue of Blackburn’s collection see p. 270]; Type [Blackburn]
Australia, Blackburn Coll. BM. 1910 : 236; Pseudolacon rufus Blackb. [Blackb.]
(BMNH).

The description is based on an unrecorded number of specimens from Roebuck
Bay sent by Mr French.

Agrypnus rugatus (Candéze) comb. n.
Tilostarsus [sic] vugatus Candéze, 1857 : 174.
Holotype. Mapacascar: 9, Madagascar; Sainot, Madag.; Janson coll. 1903 : 130;

Tilotarsus rugatus Cand. [Cand.]; Tilotarsus rugatus Cdze., type, ex coll. Deyrolle
[Janson] (BMNH).

Agrypnus rugosus (Fleutiaux) comb. n.
Adelocerva (Archontas) vugosa Fleutiaux, 1934) : 56.
LECTOTYPE (present designation). MADAGASCAR: 9, Périnet, forét céte est;
rugosa Fleut. type [Fleut.] (MNHN, Paris).

Paralectotypes. 4 ex, same locality as lectotype (MNHN, Paris). I 9, same
locality as lectotype with Fleutiaux’s label ‘Adelocera (Lacon) rugosa Fleut.’
(BMNH).

Agrypnus rusticus (Candéze) comb. n.

Tilotarsus [sic] vusticus Candéze, 18930 : 13.
Lobotarsus vusticus (Candéze) Schwarz, 1906 : 30.

The description is based on an unrecorded number of specimens from MADAGASCAR;
Nossi-Be received from M. le Dr Brancsik, of Trencsin.
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on the following specimen: g, Madagascar,
Collection Leon Fairmaire, 1906; Tylotarsus rusticus Cand. [Fleut.] (BMNH). The
metasternum and wings are much reduced in length.

Agrypnus sakaguchii (Miwa) comb. n.
Lacon sakaguchi Miwa, 1927 : 14; pl. 1, fig. 3.
Adelocera (Adelocera) sakagucht (Miwa) Ohira, 19670 : 130.

Holotype. JAPAN: 9, Loo-Choo, Okinawa-honto, collected by S. Sakaguchi.
The location of the type is not recorded. ?Taiwan Agricultural Research Institute,
Taipei.

The generic attribution is based on the description and figure.

|

—————

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 207

Agrypnus samburensis (Fleuteaux) comb. n.

Lacon samburensis Fleutiaux, 1919 : 22.
Tylotarsus (Lobitarsus [sic}) samburensis (Fleutiaux), 1935¢ : 94

LECTOTYPE (present designation). KENYA: 3, Samburu; Lacon samburensis
Fleut. type [Fleut].; Fleut. Vaj. All. Jeannel. Afr. or. Elat. p. [Fleut.] (MNHN,
Paris).

Paralectotype. 3, Samburu, 30.x to 20.xi.g6 (MNHN, Paris).

Agrypnus scaber (Candéze) comb. n.

Lacon scaber Candéze, 1857 : 53.
Lacon scabey Candéze; Neboiss, 1961 : 9.

Holotype. Mapacascar: 3g, Lafert. Madagascar; ... [illegible word] Madag.?;
285; S. scaber Cdz. type [Cand.]; Lacon scaber Cdze, Cand. type. e coll. de Laferte
[Janson] (BMNH).

Agrypnus scarrosus (Candéze) comb. n.

Lacon scarrosus Candéze, 1857 : 157.
Pyrganus scarvrosus (Candéze) Golbach, 1968 : 108.

LECTOTYPE (present designation). Merxico: ¢ Tuspan; 761; E. scarrosus
[Cand.]; Collection Chevrolat; scarrosus Cand., typ. Mon. p. 157 [Fleut.] (MNHN,
Paris). The specimen stands over a green Chevrolat collection label; Lacon scarrosus
Chevr. Cand. type Mon. 1. p. 157, no. 88, Collection Chevr. D. A. Salle.

Paralectotype. 9, Mexico, Janson coll. 1903.130; Lacon scarrosus Cand. Cand.
Type e coll. de Laferte [Janson]; scarrosus, [sic.] Mexico Chevr. [? Chevr.] (BMNH).
It is of interest to note that the lateral margins of the prothorax of the paralecto-
type are distinctly, ifirregularly crenate. (cf. Golbach’s generic diagnosis of Pyrganus),

Agrypnus schwaneri (Candéze) comb. n.

Lacon schwanert Candéze, 1874 : 73.
Adeloceva (Compsolacon) schwaneri (Candéze); Van Zwaluwenburg, 1959 : 354.

Lectotype (Van Zwaluwenburg, 1959 : 354, see p. 9). Sex undetermined.
Borneo (RNH, Leiden). It seems probable that Van Zwaluwenburg’s statement
‘[type] designated by Candéze’ indicates that the specimen bears Candéze’s deter-
mination label, possibly with the word ‘type’. Candéze’s second specimen, in the
Van Bruck collection, was lost in the bombing of Bonn (Van Zwaluwenburg, loc.
cit.).

The generic attribution is based on the description.

208 CHM. BP. VON SE AVETK

Agrypnus schwarzi (Schenkling) comb. n.

Lacon nebulosus Schwarz, 19056 : 278. [Junior primary homonym of Lacon nebulosus Candéze,
1857.]
Lacon schwarz Schenkling, 1925 : 33. [Replacement name for nebulosus Schwarz.]
Adeloceva schwarz (Schenkling) Fleutiaux, 1926 : 99.
The description is based on an unrecorded number of specimens from MapDa-
GASCAR, Cap d’Ambre.
Type-material: ?DEI, Eberswalde.
The generic attribution is based on the following specimen. MADAGASCAR:
Mont d’Ambre; nebulosus Schw., comparé au type [Fleut.] (MNHN, Paris).

Agrypnus scopulosus (Elston) comb. n.

Lacon scopulosus Elston, 1924 : 204.
Lacon scopulosus Elston; Neboiss, 1956 : 12.

The description is based on an unrecorded number of specimens from Queensland;
Coen River (W. D. Dodd) and Endeavour River.

Type-material (see p. 274): SAM, Adelaide, AM, Sidney.

Syntype examined. AUSTRALIA: g, Coen R. Q. W. D. Dodd; co-type (SAM,
Adelaide).

Agrypnus scopus (Schwarz) comb. n.

Lacon scopus Schwarz, 1902b : 200.
Adelocera scopus (Schwarz) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). NEw GuINEA: 4, N. Guinea, Stephansort;
coll. Schwarz; Typus; Lacon scopus Schw. [Schwarz] (DEI, Eberswalde).

Agrypnus scrofa (Candéze) comb. n.

Lacon scrofa Candéze, 1873 : 4.
Colaulon (Cryptolacon) scrofa (Candéze); Kishii, 1964 : 26. [Good bibliography.]

LECTOTYPE (present designation). JAPAN: g, right hand specimen of two
ononecard. Kobe, 71 [underside of card mount]; Kobe; Japan, G. Lewis 1910 : 320;
Lacon scrofa n.sp. Cdze [Lewis] (BMNH). The absence of Candéze’s determination
label is probably due to Lewis, who appears to have been in the habit of replacing
other workers’ determination labels with his own.

Paralectotypes. I g, on same card as lectotype (BMNH). 1 9, 416, 5/1/67;
Scropha [sic] Cdz. Jap. [Cand., yellow border]; collection E. Candéze; Lacon scrofa
Cd. det. E. Candéze [IRSNB curatorial label] (IRSNB, Brussels).

Candéze records that he has seen a dozen specimens, but I have been unable
to discover any more specimens collected on Lewis’s first expedition to Japan.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 209

Agrypnus scrofilia (Ohira) comb. n.
Colaulon (Cryplolacon) scrofilia Ohira, 19696 : 43.

Holotype. JAPAN: gj, Oshima Is. 28-III. 1952. H. Ohira. Ohira collection.
Paratypes. 24,39, ibidem. Ohira collection.
The generic attribution is based on the description.

Agrypnus sculptus (Candéze) comb. n.

Lacon sculptus Candéze, 1874 : 95.
Lacon sculptus Candéze; Neboiss, 1956 : 12.

The description is based on many specimens from the Paroo and Darling Rivers
obtained from de Castelnau.

Type-material: ?IRSBN, Brussels.

The generic attribution based on the following specimens, AUSTRALIA (N.S.W.):
2 3, 3 2, 2 ex, (?sex, abdomen missing), N. Holl. Paroo; Janson coll. 1903.130.
One specimen bears Gahan’s determination label ‘sculptus Cand.’. These specimens
may be part of the syntype series.

Agrypnus scutellaris (Candéze)

Lacon scutellaris Candéze, 18936 : 9.
Agrypnus scutellaris (Candéze) Nakane & Kishii, 1955 : 4.

The description is based on an unrecorded number of specimens from southern
Japan, Liu-Kiu archipelago: Oshima.

Type-material: ?IRSNB, Brussels.

Candéze states that the species bears a resemblance to binodulus and murinus,
which suggests that Nakane and Kishii’s generic attribution is fully justified.

I have been unable to find the specimens from Yokohama and the other localities
recorded by Lewis (1896 : 336).

Agrypnus scutellatus (Candéze) comb. n.

Lacon scutellatus Candéze, 1857 : III.
Archontas scutellatus (Candéze) Fleutiaux, 1947 : 283.

The description is based on an unrecorded number of specimens from ‘Des Indes —
Orientales [see p. 271] et de Java’, which Candéze received from de la Ferté Sénectére,
Mniszech and Deyrolle.

Syntype examined. JAvA: 9, Laf. Java; Janson coll. 1903.130; scutellatus
[Cand.]; Lacon scutellatus Cdze. Cand. Type e coll. de Laferté [Janson] (BMNH).
In this specimen the propleural groove mentioned in the description is very indistinct.
I have been unable to trace the specimens from the Deyrolle and Mniszech collec-
tions. If Candéze retained them they should be in the BMNH or possibly the

O

210 C2 MM. FS Von ts HAY EK

IRSNB, Brussels. If he returned them to their owners, Deyrolle’s material should
be in the BMNH (see p. 274) and Mniszech’s in the IRSNB, Brussels (see p. 278).

Agrypnus semivestitus (Elston) comb. n.

Lacon semivestitus Elston, 1927 : 352.
Adelocera (Compsolacon) semivestitus (Elston); Van Zwaluwenburg, 1966 : 298.

The description is based on an unrecorded number of specimens from N.-W.
AUSTRALIA: Wyndham (W. Crawshaw), N. Queensland: Townsville (G. F. Hill),
Groote Eylandt (N. Tyndale).

Syntypes (see p. 274). AM, Sydney, SAM. Adelaide.

The generic attribution is based on the following specimen; AUSTRALIA: 9, Cairns,
standing as semivestitus Elston (SAM, Adelaide).

Agrypnus septentrionalis (Fleutiaux) comb. n.

Adeloceva (Archontas) septentrionalis Fleutiaux, 1934b : 50.

LECTOTYPE (present designation). MADAGASCAR: 3, Diego Suarez; septen-
trionalis Fleut. type [Fleut.] (MNHN, Paris).

Paralectotypes: 3g, Diego Suarez; Alluaud; septentrionalis Fl. co-type [Fleut.].
I g, Diego S.; Museum Paris, Madagascar, coll. Perrier de la Bathie, 1906 (MNHN,
Paris).

Agrypnus serricollis (Candéze) comb. n.

Lacon serricollis Candéze, 1857 : 140; pl. 2, fig. Io.
Adelocera serricollis (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). JAvA: 9, Java; Janson coll. ex Candéze,
1903.130; Lacon serricollis Cdz. Java [Cand.]; Lacon serricollis Cand. Type ex coll.
Candéze [Gahan] (BMNH).

Paralectotypes: 1 3, Dej. Java [Janson]; Janson coll. 1903.130; Lacon serricollis
Cdze, Cand. [Janson]; serricollis [Cand.]; Agrypnus serricollis Dej. Cat. e. coll.
Dejean [Janson]. 2 9, with labels as above but without Candéze’s determination
label. 1 3, Java; Janson coll. 1903.130; Lacon serricollis Cand. ex coll. Cand.
[Gahan] (BMNH).

Agrypnus serrula (Candéze) comb. n.

Lacon serrula Candéze, 1857 : 122.
Compsolacon serrula (Candéze) Fleutiaux, 1949 : 285.

LECTOTYPE (present designation). 9, Laf. Ind. bor. Bacon [Janson]; Janson
coll. 1903.130; serrula [Cand.]; Lacon serrula Cdz. Type e coll de Laferte [Janson]
(BMNH). The specimen measures 11 mm in length compared with the published
length of Io mm.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 211

The published locality is ‘Indes-Orientales’, see p. 273. The discrepancy between
the published locality and the label is probably due to Janson, see p. 276.

Agrypnus setiger (Bates)

Lacon setiger Bates, 1886 : 338.
Agrypnus setigery (Bates) Ohira, 1954a : 4.
LECTOTYPE (present designation). TAtwan: 9, Ins. Formosa; Janson coll.

1903.130; Lacon setiger Bates Formosa [Bates]; Lacon setiger Bates Type (ex coll.
Bates [Janson]) (BMNH).

Agrypnus setosus (Schwarz) comb. n.

Lacon setosus Schwarz, 19026 : 311.
Adelocera setosus (Schwarz) Fleutiaux, 1926 : 26.

The description is based on an unrecorded number of specimens from MADAGASCAR,
Antongil Bai.

Type-material: ?DEI, Eberswalde.
The generic attribution is based on the following specimen: 9. MADAGASCAR,

Fenerive, E. Perrot; setosus Schw. comparé au type, Janv. 42 [Fleut.] (MNHN,
Paris).

Agrypnus setulosus (Candéze) comb. n.

Lacon setulosus Candéze, 1882 : 8.
Adeloceva setulosus (Candéze) Fleutiaux, 1926 : 96.

Lectotype (Van Zwaluwenburg, 1959 : 352): LESSER SUNDA ISLANDS: n.sp.
setulosus Cdz. Sumbawa Lsb. [probably Cand. with yellow border]; ex typus (IRSNB,
Brussels). Not examined.

Paralectotypes. 1 ex, standing beside lectotype (IRSNB, Brussels). Van
Zwaluwenburg does not record the labels on this specimen. 1 9, Sumbawa, Coiffs;
setulosus Cdz. nov. sp. [Cand.]; Museum Paris, Coll. R. Oberthtir, 1952. 3 4,
same locality without determination labels (MNHN, Paris).

The description is based on six specimens from Sumbawa and Flores collected
by Coiffs in the Lansberge collection. The missing specimens may now be in the
MNHN, Paris (Lansberge coll. see p. 276).

Agrypnus seyrigi (Fleutiaux) comb. n.

Adeloceva (Arvchontas) seyrigi Fleutiaux, 1934) : 56.
LECTOTYPE (present designation): MApDAscAR: g, Madagascar, Bekily. Reg.

| Sud. de VIle; seyrigi Fleut. type [Fleut.] (MNHN, Paris).

212 CZ Ma ES vow EVAWEENKK

Paralectotypes. 49, 2 ex, locality as lectotype. Two specimens bear Fleutiaux’s
determination label (MNHN, Paris).

Agrypnus silvatica (Fleutiaux) comb. n.
Adelocerva sylvatica Fleutiaux, 1932¢ : 153.

LECTOTYPE (present designation). MADAGAsCcAR: 92, Madagascar, Lavauden;
Forét de Tempina, Est Madag. Sud Tamatave; Adelocera (Lacon) silvatica Fleut.
type [Fleut.] (MNHN, Paris).

Agrypnus similis (Fleutiaux) comb. n.
Tylotarsus similis Fleutiaux, 1934) : 60.

LECTOTYPE (present designation). MADAGASCAR: 2, Fort Dauphin, Tylotarsus
similis Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 7 ex., Fort Dauphin. 2 ex., Madagascar Sud, Foréts Nord
Ft. Dauphin; Museum Paris, Madagascar, Coll. Ch. Alluaud 1904. One specimen
bears Fleutiaux’s determination label. 4 ex., Madagascar Sud, Andrahomana,
Alluaud Igo00. xi. One specimen bears Fleutiaux’s determination label. I ex.,
Madagascar Sud, Vallée d’Ambolo, Alluaud 1900. 1 ex, Museum Paris, Madagascar,
Grandidier 1851.91. 1 ex, Museum Paris, Madagascar, Tananerive, Waterlot,
I9g16. 1 ex, Museum Paris, Madagascar, Province de Fort Dauphin, Antanimora,
R. Decary, 1926. 8 ex, Museum Paris, Madagascar, Collection Perrier de la Bathie,
1906 (MNHN, Paris). The paralectotypes from “entre Soanierana et Foulepointe
(J. Descarpentries)’ have not been located.

Agrypnus simplex (Candéze) comb. n.

Tilotarsus [sic] simplex Candéze, 1874 : IIo.
Lobotarsus simplex (Candéze) Schwarz, 18984 : 131.

LECTOTYPE (present designation): GABON: g, Gabon; Janson Coll. 1903 : 130;
simplex Cdz. type [Cand.]; Tilotarsus simplex Cdze, type [Janson] (BMNH).

Paralectotype: 3, Gabon, Janson coll. 1903 : 130; Tilotarsus simplex Cdze [ Janson]
(BMNH).

Agrypnus sinensis (Candéze) comb. n.

Lacon sinensis Candéze, 1857 : 139.
Lacon massiei Fleutiaux, 1895a : 685. [Synonymized by Fleutiaux, 1918d : 192.]
Compsolacon sinensis (Candéze) Fleutiaux, 1947 : 285.

Lacon sinensis Candéze. LECTOTYPE (present designation). CHINA: 2,
261; China, Dubois; sinensis Cand. type Mon. [Fleut.]._ The specimen stands over
a yellow Chevrolat label: Lacon sinensis Cand., type. Mon. 1. 139. 67. China, D.
Dubois (MNHN, Paris).

————

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 213

The description is based on an unrecorded number of specimens from China in
the Dohrn and Chevrolat Collections. Fleutiaux (1g18d: 192) states that he
possesses the type from the Chevrolat collection, but does not give sufficient
information to identify the specimen. The Dohrn material may be in the
IZPAN, Warsaw (see p. 274).

Lacon massiet Fleutiaux. LECTOTYPE (present designation). Laos: 4,
Luang Prabang, Massie; Lacon massiei Fleut., type [Fleut.]; Fleut. Ann. Soc.
ent. Fr. 1894 : 685 [Fleut.]; sinensis Cand. [Fleut.] (MNHN, Paris).

Agrypnus singularis (Fleutiaux) comb. n.

Lacon singularis Fleutiaux, 1919 : 21.
Adelocera singularis (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). TANZANIA: g, Lindi, Afrique orient.
Allemande, collection Le Moult; Lacon singularis Fleut., type [Fleut.]; Fleut.
Voy. All. Jeann. Afr. or. Elat, p. [Fleut.] (MNHN, Paris).

Agrypnus sinuatus (Candéze) comb. n.

Lacon sinuatus Candéze, 1857 : 102; pl. 1, fig. 16.
Adelocera sinuatus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MapaGascar: 9, Madagascar; Janson
coll. 1903 : 130; Lacon sinuatus Cdze, Type (e coll. de Mniszech) [Janson] (BMNH).
The absence of Candéze’s determination label is probably due to Janson (see p. 276).
The BMNH manuscript catalogue of the Janson collection records that it contains
the type of sinuatus Cand. from the Mniszech collection.

Lacon sjostedtt Schwarz, 1908a : 57.

:
Agrypnus sjostedti (Schwarz) comb. n.
| Lobotarsus sjostedti (Schwarz) Schenkling, 1925 : 39.

LECTOTYPE (present designation). TANZANIA: 2, Mombo, Sjostedt; Usambara;
juni; coll. Schwarz; Typus; sjostedti Schw. [Schw.]; Tylotarsus nec Lacon [Fleut.]
(DEI, Eberswalde).

) Paralectotype: 2, Mombo, Sjostedt; typ.; Sjostedti; Schw. O. Schw. det. 1907;
typus (NR, Stockholm).

Agrypnus socius (Candéze) comb. n.
| Lacon socius Candéze, 1874 : 87.
| Adelocera socius (Candéze) Fleutiaux, 1926 : 96.

| LECTOTYPE (present designation). AUSTRALIA: g, Cape York; Janson coll.
1903 : 130; L. socius type [Cand.]; Lacon socius Cand., type Cape York [Gahan]
(BMNH).

214 C. M. F. von HAYEK

Agrypnus soleatus (Candéze) comb. n.

Tilotarsus (sic) soleatus Candéze, 1857 : 176.
Lobotarsus soleatus (Candéze) Schwarz, 1906 : 30.

Holotype. GuINEA: 9, Tylotarsus soleatus mihi [Cand.]; Tilotarsus soleatus
Cdze, type ex coll. de Laferté [Janson] (BMNH).

Agrypnus soricinus (Candéze) comb. n.

Lacon sorvicinus Candéze, 1857 : 133; pl. 2, fig. 11.
Adelocera sovicinus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MADAGASCAR: J, Laf. Madagascar; Janson
collection 1903 : 130; Lacon soricinus Cand. type [Cand.]; Lacon soricinus Cdze,
Cand., Type e coll. de Laferté [Janson] (BMNH).

Agrypnus spinifer (Candéze) comb. n.
Tilotarsus [sic] spinifer Candéze, 1889 : 74 (8).

The description is based on an unrecorded number of specimens from MADAGAS-
CAR. I would expect this material to be in the IRSNB, Brussels (see p. 273) but
Fleutiaux (1899 : 222) comments that the specimen in the MNHN, Paris labelled
Tilotarsus spinipes (see below) is probably the type, “spinipes’ being a lapsus calami.
Examination of any material extant in the IRSNB, Brussels will show whether
Fleutiaux was correct in his surmise.

The generic attribution is based on the following specimen: ?¢, (genitalia missing),
MADAGASCAR, Sainte Marie Cloué, 1847 Museum Paris. Tylotarsus spinipes sp.n.
89 [Cand.]; spinifer Elat. nouv. IV [Fleut.] (MNHN, Paris).

Agrypnus spissicollis (Candéze) comb. n.
Tilotarsus [sic] spissicollis Candéze, 18936 : 12.
Lobotarsus spissicollis (Candéze) Schenkling, 1925 : 39.
The description is based on an unrecorded number of specimens from
Antananarivo [MADAGASCAR].
Type-material: ?IRSNB, Brussels.
The generic attribution is based on the following specimen: 9, MADAGASCAR

Miarinarivo; spissicollis Cand. comparé au type [Fleut.] (MNHN, Paris). The
metasternum is much reduced and the wings entirely absent.

Agrypnus squalescens (Fairmaire) comb. n.

Tilotarsus [sic] squalescens Fairmiare, 1871 : 39.

The description is based on an unrecorded number of specimens collected in
MADAGASCAR by Ch. Coquerel.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 215

Type-material: ?IRSNB, Brussels.

The generic attribution is based on a note in the Fleutiaux collection (MNHN,
Paris) ‘squalescens Fairm, type collection Cand., Musée de Bruxelles, resemble
a rusticus Cand., plus etroit, pronotum plus long’.

Agrypnus squalida (Fleutiaux) comb. n.
Adelocera squalida Fleutiaux, 1927 : 76.

LECTOTYPE (present designation). CAmBopIA: 3, Camboge, Kampong Thon,
Vitalis; Adelocera (Lacon) squalida Fleut., type [Fleut.] (MNHN, Paris).

Agrypnus squameus (Szombathy) comb. n.

Adelocera squamea Szombathy, 1909 : 118; fig. 1.
Adelocera squamea var. obscura Szombathy, loc. cit.

LECTOTYPE (present designation). E. Arrica (position unknown): 9, Afr.
or. Katona; Mto-ja-Kifarou; Holotypus 1909 Adelocerasquamea Szombathy[Kaszab]
(TM, Budapest).

Paralectotype. 9, locality as lectotype, labelled ‘Paratype Adelocera squamea
Szombathy’ by Kaszab. (TM, Budapest).

I have also examined two @ from the same locality labelled as the
paratype of Adelocera squamea var. obscura Szombathy by Kaszab. All four
specimens agree well with the descriptions and despite the absence of Szombathy’s
determination labels I believe that they form the original syntype-series.

Agrypnus stictus (Candéze) comb. n.

Lacon stictus Candéze, 1895b : 56.
Adelocera stictus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens collected in
Madagascar, Diego-Suarez by Alluaud.
Type-material: P>MNHN, Paris. Not located, 1971.

The generic attribution is based on the following specimen; MADAGASCAR: 4, Ste
Marie de Madagascar; stictus Cand., comparé au type Xr. 43 [Fleut.] (BMNH).

Agrypnus stigmosus (Elston) comb. n.

Lacon stigmosus Elston, 1927 : 352.
Lacon stigmosus Elston; Neboiss, 1956 : 13.

The description is based on an unrecorded number of specimens from N.W.
Australia: Derby and N. Queensland: Normanton (R. Kemp).

Type-material (see p. 274): SAM, Adelaide, AM, Sydney, NM, Victoria.

216 Cee GBs WiONe vEARYGE KK

The generic attribution is based on two syntypes; AUSTRALIA: 2 9, Queensland;
Lacon stigmosus Elston Cotype [?Elston] (SAM, Adelaide).

Agrypnus stricticollis (Fairmaire) comb. n.

Lacon stricticollis Fairmaire, 1881 : 266.
Adeloceva stricticollis (Fairmaire) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from the Viti
Islands [Fiji]. Fairmaire records that he received material for his work on the
fauna of these islands from Godeffroy of Hamburg, but does not state whether he
retained any or all of it. Any material retained by Fairmaire should be in the
IRSNB, Brussels or MNHN, Paris (see history of the Fairmaire collection,
p- 275) but up to the present time it has not been located. The MNHN, Paris contains
a specimen from Ins. Fidji (Watkins) determined by Fleutiaux with a note ‘comp.
au type Mus. Brux. Nov. 32’. Godeffroy (Horn & Kahle, 1935 : 91) did not retain
any duplicate material in his museum, which was broken up in 1885, part being
acquired by the Zoological Museum, Hamburg.

The generic attribution is based on the following specimens which may be part
of the syntype-series. Fit: 2 g, Viti, 81:50 [BMNH registration number =
purchased of Godeffroy Museum, Hamburg, 1881]. One specimen bears a label
‘Lacon sticticollis [sic] Fairm.’ in an unknown hand (BMNH).

Agrypnus suarezi nom. n.

Lobotarsus maculatus Schwarz, 19050 : 280.
Agrypnus maculatus (Schwarz) comb. n. [Junior secondary homonym of Agrypnus maculatus

Macleay, 1872.]

The description is based on an unrecorded number of specimens from Madagascar,
Diego Suarez.

Type-material: >DEI, Eberswalde.

The generic attribution is based on the following. specimen; MADAGASCAR: Mt.
de Ambre, Nov.; Lobotarsus maculatus Schw. comp. au type [Fleut] (MNHN,
Paris).

Agrypnus subcarinulatus (Schwarz) comb. n.

Lacon subcarinulatus Schwarz, 1908a : 82.
Adelocera subcarinulatus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from MADA-
GASCAR, Mont d’Ambre.
Type-material: ?>DEI, Eberswalde.

The generic attribution is based on Schwarz’s comment that the species bears
a resemblance to sulcicollis Schwarz (see p. 219). A note in the Fleutiaux collection

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 217

(MNHN, Paris) states ‘subcarinulatus Schw. vu type = crenatus Klug’. Fleutiaux
did not publish the synonymy.

Agrypnus subcervinus (Fleutiaux) comb. n.

Lacon subcervinus Fleutiaux, 1916 : 219.
Adelocera subcervinus (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). PHILIPPINES: 9, Luzon, Mont. Trimay;
Lacon subcervinus Fleut., type [Fleut] (MNHN, Paris).

Agrypnus subcylindricus (Schwarz) comb. n.

Lacon subcylindricus Schwarz, 1908c : 93.
Adelocera subcylindricus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from MaApa-
GASCAR, Mont d’Ambre.
Type-material: ?>DEI, Eberswalde.

The generic attribution is based on the following specimen: MADAGASCAR: 9,
Diego Suarez; subcylindricus Schw. comp. au type [Fleut.] (MNHN, Paris).

Agrypnus submarmoratus (Elston) comb. n.

Lacon submarmoratus Elston, 1924 : 205.
Lacon submarmoratus Elston; Neboiss, 1956 : 13.

The description is based on an unrecorded number of specimens from [AUSTRALIA]
Queensland: Cairns (A. H. Lea).

Type-material (see p. 274): SAM, Adelaide, AM, Sydney, NM, Victoria.

The generic attribution is based on the following specimens; AUSTRALIA: I 4,

2 9, from the type-locality standing as co-types of submarmoratus Elston (SAM,
Adelaide).

Agrypnus subocellatus (Candéze) comb. n.

Lacon subocellatus Candéze, 1882 : 6.
Adelocera subocellatus (Candéze) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Nossi
Bé [MADAGASCAR].
Type-material: ?IRSNB, Brussels, but see below.

The generic attribution is based on the following specimen, MADAGASCAR: g,
Nossi Bé, recu du Boyer; Lacon subocellatus Cdz. type [Cand.] (MNHN, Paris).
This specimen may be part of the syntype series.

218 CM, EF. Von hank

Agrypnus suboculatus (Candéze) comb. n.

Tilotarsus [sic] suboculatus Candéze, 1882 : 13.
Lobotarsus suboculatus (Candéze) Schwarz, 1906 : 30.

The description is based on an unrecorded number of specimens from
Antananarivo [MADAGASCAR].
Type-material: ?>IRSNB, Brussels.

The generic attribution is based on the following specimen: 9, MADAGASCAR,
Vohemar, Coll. Le Moult; Tylotarsus suboculatus Cand. [Fleut] (BMNH).

Agrypnus subsericeus (Candéze) comb. n.

Lacon subsericeus Candéze, 1878a : 102.
Adelocera subsericeus (Candéze) Fleutiaux, 1926 : 96.

Holotype. NEw Guinea, Hatam (Beccari) >MCSN, Genoa.
The generic attribution is based on the description.

Agrypnus subserratus (Quedenfeldt) comb. n.

Tylotarsus subservratus Quedenfeldt, 1886 : 24; pl. I, figs 14, 14a & b.
Lobotarsus subservatus (Quedenfeldt) Schwarz, 1906 : 30.

LECTOTYPE (present designation). , Tylotarsus subserratus Qdf. Qgo [Qued.];
Ex Museo Quedenfeldt (MNHN, Paris).

The published locality is Quango [= Kwango or Cuango] River, collected by
v. Mechow. The exact location is unknown.

Agrypnus substriatus (Fleutiaux) comb. n.
A deloceva (Archontas) substriata Fleutiaux, 1934) : 55.
LECTOTYPE (present designation). MADAGASCAR: dg, Madagascar, Forét

Tanala, Alluaud, 1901; Museum Paris, Madagascar, coll. Alluaud, 1904; substriata
Fleut., type [Fleut] (MNHN, Paris).

Paralectotypes. 1 3, I 9, same locality and collection as lectotype but without
determination labels. 19, Museum Paris, Madagascar, Mahasinjo, Coll. J. Chatanay,
Ig14. 19, Nanisana: Madagascar, Tananarive (MNHN, Paris).

Agrypnus subtuberculatus (Schwarz) comb. n.
Lobotarsus subtuberculatus Schwarz, 1898a : 132.

The description is based on an unrecorded number of specimens from Kameroun
[CAMEROUN], collected by Conrad and submitted by Kraatz.

Type-material: ?>DEI, Eberswalde.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 219

The generic attribution is based on the following specimen. EQUATORIAL GUINEA:
Benito, Congo franc.; Lobotarsus subtuberculatus Schw. [Schwarz] (MNHN, Paris).

Agrypnus suillus (Candéze) comb. n.

Lacon suillus Candéze, 1857 : 159.
Pyrganus suillus (Candéze) Golbach, 1968 : 1098.

Holotype. Mexico: Dej. Mexico; Janson coll. 1903.130; Lacon suillus (De}j.)
Cand. Cand. Type [Janson]; Agrypnus suillus Dej. coll. e coll. Dej. [Janson] (BMNH).

The abdomen is missing. The absence of Dejean and Candéze determination
labels is probably due to Janson (see p. 276).

Agrypnus sulcicollis (Schwarz) comb. n.

Lacon sulcicollis Schwarz, 1908a : 73.
Adelocera sulcicollis (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from Mapa-
GASCAR, Antananarivo.

Type-material: ?>DEI, Eberswalde.

The generic attribution based on the following specimen; MADAGASCAR: ,
Antongil; sulcicollis Schw., comparé [Fleut.] (MNHN, Paris).

: Agrypnus taciturnus (Candéze) comb. n.

| Lacon taciturnus Candéze, 1874 : 60.
Adelocera (Adelocera) taciturna (Candéze) ; Ohira, 19676 : 103.

The description is based on an unrecorded number of specimens from Laos in
the Janson and Candéze collections.

Syntypes examined. 1 3, Muhot, Laos; Janson coll. 1903.130; Lacon taciturnus
Cand. Laos (Mouhot) [Gahan]. 1 4, Muhot, Laos; Janson coll. 1903.130; L.
taciturnus Cdz. [Cand.]; Lacon taciturnus Cand., Cotype, Laos [Gahan] (BMNH).
The IRSNB, Brussels may contain additional syntype specimens, see notes on the
Candéze collection, p. 000.

Agrypnus colonicus (Candéze) may be conspecific with this species, see p. I4I.

Miwa (1934 : 168) placed Paralacon koshunensis Miwa (1929 : 233) in synonymy
with ¢aciturna Candéze and Ohira (1969c : 28) treats kosunensis [sic] as a subspecies
of taciturna. Clarification of the status of koshunensis must be postponed until the

| type-material has been located and examined.

Lacon tactus Candéze, 1874 : 67.
Adelocera tactus (Candéze) Fleutiaux, 1926 : 96.

| Agrypnus tactus (Candéze) comb. n.

220 C. M. F. von HAYEK

LECTOTYPE (present designation). THAILAND. ¢, Janson coll. 1903 : 130;
tactus Cdz, type [Cand.]; Lacon tactus Cand., type. Siam [Gahan] (BMNH).

Agrypnus taiwanus (Miwa) comb. n.
Lacon taiwanus Miwa, 1927 : 13; pl. I, fig. 2.
Syntypes. Formosa: 4 ex., Baibara (A. Saito), Koshun (J. Sonan), Horisha and

Arisan (Dr S. Matsamura). The location of the material is not recorded. ?Taiwan
Agricultural Research Institute, Taipei.

The generic attribution is based on the description.

Agrypnus tectus (Fleutiaux) comb. n.
Tylotarsus (Lobitarsus [sic]) tectus Fleutiaux, 1935¢ : 92.

LECTOTYPE (present designation). KENya: Turner —- McArthur, Lower Tana —
Sabaki, April-May 1932; Tylotarsus tectus Fleut. [Fleut.] (MNHN, Paris).

Paralectotype. 9, same locality as lectotype and bearing Fleutiaux’s determina-
tion label (NM, Nairobi).

Agrypnus tellini (Fleutiaux) comb. n.

Lacon tellini Fleutiaux, 1903¢ : 250.
Adelocera tellini (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). Eruiopia: 3, Ghinda; Erithrée Tellini;
Lacon tellini Fleut., type [Fleut.]; Bull. Soc. Ent. Fr. 1903 p. 290 [Fleut.] (MNHN,
Paris).

Agrypnus terminata (Fleutiaux) comb. n.
Adelocera (Archontas) terminata Fleutiaux, 1934) : 53.

LECTOTYPE (present designation) 9, MApAGAscAR, Maevetanana; terminata
Fleut., type [Fleut.] (MNHN, Paris).

Agrypnus testaceus (Schwarz) comb. n.
Centrostethus testaceus Schwarz, 1908d : Io.
The description is based on an unrecorded number of specimens from Madagascar,
Antananarivo.
Type-material: >DEI, Eberswalde.

The generic attribution is based on the following specimen: 3, MADAGASCAR:
Marvansetra; Decembre; Centrostethus testaceus Schwarz comparé [Fleut.] (MNHN,
Paris).

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 221

Agrypnus thomasi nom. n.

Homeolacon gracilis Blackburn, 1890 : 91.

Agrypnus gracilis (Blackburn) comb. n. [Junior secondary homonym of Agrypnus gracilis
(Candéze, 1874).]

LECTOTYPE (present designation). AUSTRALIA: g, 2800, T., N.T. [on card
mount, see p. 270]; Australia, Blackburn coll. BM 1910.236; Homeolacon gracilis
Blackb. [Blackb.] (BMNH). The published locality is ‘Northern Territory’.

Paralectotypes. IRSNB, Brussels and SAM, Adelaide according to Neboiss
(1956 : 15 and 1961: 10). Not examined.

Agrypnus thibetanus (Reitter) stat. nov., comb. nov.

Compsolacon crenicollis var. thibetanus Reitter, 1913 : 658.

The description is based on numerous specimens from Thibet: Poo, West Himalaya
submitted by Spichal.

Type-material: ?>TM, Budapest.

The generic attribution is based on the following specimens: CHINA: I 9, Poo,
West. Hym. Coll. Spichal; thibetanus Rtt [?Reitter]; lapideus Cand. [Fleut.] 1 3,
same locality; Compsolacon thibetanus Rtt [Fleut.] (MNHN, Paris). These speci-
mens are probably part of the syntype-series.

These specimens are not conspecific with the specimens from the Caucasus stand-
ing as crenicollis in the BMNH and MNHN, Paris.

Agrypnus tigrinus (Fleutiaux) comb. n.
Lacon tigrinus Fleutiaux, 1919 : 23.
Adelocera tigrinus (Fleutiaux) Fleutiaux, 1926 : 96.
LECTOTYPE (present designation). TANZANIA: g, Dar es Salaam; Lacon

tigrinus Fleut., type [Fleut.]; Fleut. Voy. All. & Jeann. Afr. or Elat. p. [Fleut.]
(MNHN, Paris).

Agrypnus tonkinensis (Fleutiaux) comb. n.
Adelocera tonkinensis Fleutiaux, 1927 : 81.
Colaulon (Sagojyo) tonkinensis (Fleutiaux) Kishii, 1964 : 34.
LECTOTYPE (present designation). NorTH VIETNAM: 9, Tonkin, Lac Tho,
Hoa Binh. A. de Cooman; tonkinensis Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 35 ex, same locality labels as lectotype; 4 specimens bear
Fleutiaux’s determination label (MNHN, Paris). 1 3, 7 9, same locality labels
as lectotype; 4 2 specimens bear Fleutiaux’s determination labels (BMNH).

The material recorded from Hoa Binh (L. Dupont) by Fleutiaux has not been

| located.

222 Gi Ma ES von HAYEK

Agrypnus torrefactus (Candéze) comb. n.

Lacon torvefactus Candéze, 1857 : 128.
Adelocera torrefactus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). 9, Deyr. India [Janson]; 283; Janson
coll. 1903.130; Lacon torrefactus Cdze Type e coll. Deyrolle (BMNH). [Janson]
(BMNH).

Paralectotype. 1 9, Deyr. India; Lacon torrefactus Cdze. Cand. e. coll. Deyrolle
[Janson] (BMNH). The published locality is ‘Indes-Orientales’. see p. 271.

Agrypnus torresi (Candéze) comb. n.

Lacon torvest Candéze, 1897 : Io.
Lacon torrest Candéze; Neboiss, 1956 : 13.

LECTOTYPE (present designation). AUSTRALIA: g, Thursday Isd.; nsp.
1894, torresi Cand. Thursday Island [Cand.]; Lacon torresi Cand. det. E. Candeze
[IRSNB curatorial label]; Collection E. Candéze (IRSNB, Brussels).

The published locality is Cape York. Thursday Island lies about 20 miles off
Cape York. The specimen agrees well with the description, except that is it 14-5 mm
long compared with the published length of 13 mm.

Agrypnus tostus (Candéze) comb. n.

Lacon tostus Candéze, 1857 : 129.
Adelocera tostus (Candéze) Fleutiaux, 1926 : 98.

Holotype. 3, Deyr. India [Janson]; 373; Janson coll. 1903.130; L. tostus [Cand.]
Lacon tostus Cdze, type e coll. Deyrolle [Janson] (BMNH).
The published locality is ‘Indes-Orientales’, see p. 271.

Agrypnus transversicollis (Fleutiaux) comb. n.

Adeloceva tvansversicollis Fleutiaux, 1927 : 80.

LECTOTYPE (present designation). Laos: g, Xieng-Khouang; 12.5.9 [sic,
?1919]; Adelocera transversicollis Fleut., type [Fleut.] (MNHN, Paris).

Paralectotypes. 3 3, I 9, same locality as the lectotype, one bears Fleutiaux’s
determination label (MNHN, Paris).

I have not found the syntype specimens from Laos: Tranninh, juin (Vitals
de Salvaza) and ANNAM: Kieng-Trap, mai (Vatalis de Salvaza).

Agrypnus transversus (Candéze) comb. n.

Lacon transversus Candéze, 1857 : 123.
Adelocera transversus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). 9 Laf. Ind. bor. Bacon [Janson]; Janson

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 223

coll. 1903.130; transversus [Cand.] Lacon transversus Cdze. Cand. Type e coll.
de Laferté [Janson] (BMNH).

The description is based on material from ‘Indes-orientales’ (see p. 271) in the
de la Ferté and Deyrolle collections. I have been unable to locate the specimens
from the Deyrolle collection.

Agrypnus tripartitus (Candéze) comb. n.

Lacon tripartitus Candéze, 1874 : 54.
Adelocera tripartitus (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). MapaGaAscar: 9, Lacon n.sp. [Janson];
tripartitus Cdz [Cand.] Lacon tripartitus type Cand. [Gahan] (BMNH).

Agrypnus truncatus (Herbst) comb. n.

Elater truncatus Herbst, 1806 : 95; pl. 166, fig. 2.

Lacon cylindricus Candéze, 1857 : 121. [Synonymized by Candéze, 18gIc : 20.]
[Lacon melancholicus Candéze; Candéze, 1892 : 484. Misidentification. }

A deloceva truncatus (Candéze) Fleutiaux, 1926 : 96.

Elater truncatus Herbst. LECTOTYPE (present designation). ¢, Hbst.;
16121 [MNHU, Berlin, museum catalogue number, refers to information on following
label]; India or. Westerm.; truncatus n. Elat. truncatus Hbst., cylindricus Cand.,
Ind. or. Hbst. [Gerstaecker, yellow paper] (NMHU, Berlin).

Paratypes. 2,19, India or. Westerm. No. 16121 (NMHU, Berlin).

Lacon cylindricus Candéze. LECTOTYPE (present designation). g, Laf.
N. India, Bacon [Janson]; Janson coll. 1903.130; 42; Lacon cylindricus Cdze,
Cand., e coll. de Laferté [ Janson] (BMNH).

Paralectotype J, labels as lectotype, but bearing the number 43 (BMNH).

The published locality is ‘des Indes-orientales’, (see p. 271). Janson (see
p. 276) is probably responsible for the discrepancy between the published locality
and the labels and also for the absence of Candéze’s determination label.

Schenkling (1925 : 27) lists jurulosus Candéze as a synonym of truncatus Herbst
but gives no reason for doing so. I have been unable to trace the type-material
of jurulosus (see p. 172).

Other material examined. Inp1A: Barwai, Chota Nagpore, 1892, 19. Nagpore,
Io 2 (IRSNB, Brussels). None of the specimens bear Candéze’s determination
label but all carry the IRSNB curatorial label ‘Lacon melancholicus Cd.’. They
are probably the specimens recorded by Candéze (1892 : 484).

Agrypnus truquii (Candéze) comb. n.

Lacon truquit Candéze, 1874 : 101.
Pyrganus truquii (Candéze) Golbach, 1968 : 198.

224 C. M. F. von HAYEK

LECTOTYPE (present designation). MExiIco: $ 22832 [Fry manuscript cata-
logue no. = Type, Mexico, San Carlos, Truqui]; Truqui, Mexico; Fry coll. 1905 : roo,
L. truquii Cdz, type [Cand.]; Lacon truquii Cand., Type, Mexico [Fry] (BMNH).

Paralectotype. 4, Oaxaca, Mexico, Salle; Lacon truquii Cand. Salle coll. 1424
{unknown manuscript, the number refers to the locality recorded in Salle’s
manuscript catalogue] B.C.A. Col. 111 (1), Lacon truquii Cand. [Champ.] (BMNH).

Agrypnus tuberculatus (Candéze) comb. n.

Tilotarsus [sic] tuberculatus Candéze, 1874 : 109.
Lobotarsus tuberculatus (Candéze) Schwarz, 1906 : 30.

LECTOTYPE (present designation). ANGOLA: g, Benguela; T. tuberculatus
Cdz, n.sp., type [Cand.]; Tilotarsus tuberculatus Cdze, type [Janson] (BMNH).

Agrypnus turbatus (Candéze) comb. n.

Lacon turbatus Candéze, 1893¢ : 169.
Adelocera turbatus (Candéze) Fleutiaux, 1926 : 98.

The description is based on two specimens of unequal size (10-12 mm) from
Belgaum, in the state of Bombay [INDIA] submitted by Andrewes. The BMNH
possesses two female specimens from this locality. The paralectotype recorded
below measures 9-5 mm in length. The other from the Janson collection bears
only Jansons label ‘L.n.sp? allied to torrefactus Cand.’ As it is also 9-5 mm long,
it cannot be the second syntype. The male specimen in the IRSNB, Brussels
carries the locality Kanara [Madras State]. However, the fact that Candéze’s
determination label bears the words ‘Bombay’ and ‘And.’[rewes] suggests that
either he overlooked the locality label or that it was not there at the time. Since
the specimen is 12-5 mm long and agrees very well with the description, I have no
hesitation in accepting it as the second syntype.

LECTOTYPE (present designation). INnprA: g, Kanara [probably erroneous,
see above] nov. sp. 1893, turbatus Cand. Bombay, And. [Cand., yellow border];
Collection E. Candéze; Lacon turbatus Cd. det. E. Candéze [IRSNB] curatorial
label] Ex typis. (IRSNB, Brussels).

Paralectotype. Inp1A: 9, Belgaum; 272; co-type; Lacon turbatus Cd. [Andrewes];
Andrewes Bequest, BM. 1922-221 (BMNH).

Agrypnus turbidus (Germar) comb. n.

Lacon turbidus Germar, 1840 : 263.
Adelocera turbidus (Germar) Fleutiaux, 1926 : 96.

The description appears to be based on specimens bearing the manuscript name
Agrypnus turbidus received from Dupont (Germar, 1840 : 440). Candéze (1857 : 99)
states that he received (presumably on loan) Germar’s ‘exemplaire typique’ from

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 225

the Berlin Natural History Museum. However the two specimens in the NMHU,
Berlin standing beside Gerstaecker’s blue label ‘turbidus Germ. Cand. Madag.
Goud.’ are 18 mm and 19 mm long compared with the published length of 7 lines
(815mm). The discrepancy in the length and the fact that the specimens came
from Goudot makes it impossible to accept them as part of the syntype-series.
The specimens are conspecific with the lectotype. The DEI, Eberswalde does not
possess any specimens determined by Germar. For the history of the Germar
collection see p. 275.

LECTOTYPE (present designation). MADAGASCAR: g, turbidus Dup. nodifer
Kl.? Madagascar [Germar]; Janson coll. ex Schaum, 1903 : 130 (BMNH).

Agrypnus turkistanicus (Schwarz) comb. n.

Lacon turkistanicus Schwarz, 1902b : 198.
Adeloceva turkistanicus (Schwarz) Fleutiaux, 1926 : 96.

The description is based on an unrecorded number of specimens from TURKISTAN,
Buchara.

Type-material: ?IRSNB, Eberswalde.

The generic attribution is based on the following specimen: 9, locality illegible,
determined by Dolin (BMNH).

Agrypnus tuspanensis (Candéze) comb. n.

Lacon tuspanensis Candéze, 1857 : 157.
: Pyrganus tuspanensis (Candéze) Golbach, 1968 : 198.

| LECTOTYPE (present designation). MExico: 9, type; L. tuspanensis [Cand.]
Collection Chevrolat; tuspanensis Cand., type Mon. p. 157 [Fleut]. The specimen
stands over a green Chevrolat label: Lacon tuspanensis Cand., type Mon. I. p. 157,
_ 89 transversus Chevr. Mexico, Tuspan. D. A. Sallé (MNHN, Paris).

Paralectotypes. 1 ¢ [not 9 as labelled], Laf. Mexico; Janson coll. 1903.130;

_ Lacon tuspanensis Cdze, Cand. Type e coll. de Laferté [Janson]. 1 9 [not gas

labelled], Laf. Mexico; Janson coll. 1903. 130; Lacon tuspanensis Cadze, Cand. e
coll. de Laferté [Janson]; transversus Chevr. Mexico [? Chevr] (BMNH).

The wrong sex signs were probably attached by Champion (see p. 9).

Agrypnus unicolor (Hope) comb. n.
Mecynocanthus unicolor Hope, 1838 : 53.

The description is based on a single specimen in Resin Animé in ‘Mr Strong’s
| superb collection’. The locality is given as ‘India oriental’.

I have been unable to find this material. It is not in the Hope collection in
_ Strong’s collection of shells and fine specimens of amber containing

|

226 CoM. ES von SEA YE

insects was sold by auction by J. C. Stevens on 6th April 1891 (Nature, April, 1881:
CLXXI and Atheneum, April, 1891: 247). Through the kindness of Mr D. B.
Janson I have been able to examine a sale catalogue annotated by his grandfather.
Lots 116 to 121 made up of ‘Specimens of Amber containing insects of various
orders’ have a pencil note “gum copal”’ beside them. Four lots were purchased
by Sowerby and two by Fulton. As these gentlemen were dealers it must be assumed
that the specimens were sold privately, and are now lost. The figure depicts an
individual with drawn out and sharply pointed anterior angles of the prothorax.
This feature does not occur in any known Indian species but is characteristic of a
number of Madagascan species. Champion (1916 : 245) suggested that Hope may
have been mistaken in the origin of his resin and that wnicolor was a pallid form of
the Madagascan Centrostethus cuspidatus (Klug). This may well be the case as
Hope (loc.cit.: 48, footnote) records that he also received Animé from Madagascar.

The generic attribution is based on the close resemblance of the figure to Agrypnus
cuspidatus (Klug).

Agrypnus unicus (Fleutiaux) comb. n.

Lacon unicus Fleutiaux, 1919 : 21.
Adelocera unicus (Fleutiaux) Fleutiaux, 1926 : 96.

Holotype. TANZANIA: 3, Afrique orientale allemande, Kilimandjaro versant
sud-est Alluaud & Jeannel; Prairies alpine du Bismarkhiigel (alt. 2600-2800,
mars—avril 1912, st. 70); Lacon unicus Fleut., type (MNHN, Paris).

Agrypnus ursulus (Candéze) comb. n.

Lacon ursulus Candéze, 1857 : 147.
Lacon ursulus Candéze; Neboiss, 1956 : 13.

LECTOTYPE (present designation). AUSTRALIA: 9, Laf. Sydney, N. Holl.
ursulus Cdz, type [Cand.]; Janson coll. 1903.130; Lacon ursulus Cdze, Cand. type
e coll. de Laferté [Janson] (BMNH).

The metasternum and wings are much reduced in length.

Agrypnus vandepolli (Candéze) sp. rev., comb. n.
Lacon Van-de-Polli Candéze, 18876 : 285.

LECTOTYPE (present designation). AUSTRALIA: 9, Normantown, Queensland
(Ch. French); Lacon Van de Polli Cand. Type [? Van de Poll]; Collection Neervoort
Van de Poll, type de Candéze [Fleut.] (MNHN, Paris).

Neboiss (1965 : 13), following Schenkling (1925 ; 30), lists vandepollt as a synonym
of variabilis Candéze. Comparison of the descriptions indicates that the synonymy
is not justified. L. vandepolli possesses golden scales and deep tarsal grooves on the
propleurae, characteristics which do not occur in variabilis. I believe that the

SSS SS ee

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 227

synonymy in the Schenkling catalogue is due to a printers error, the name variabilis
being indented and initalics. Fleutiaux (1926 : 98) published a correction indicating
that variabilis is a good species but this appears to have been overlooked by Neboiss.

Agrypnus variabilis (Candéze) comb. n.

Lacon variabilis Candéze, 1857 : 148.
Lacon variabilis Candéze; Neboiss, 1956 : 13.

LECTOTYPE (present designation). AUSTRALIA: 9, Janson coll. ex Candéze,
1903 : 130; Lacon variabilis Cdz. Austr. [Cand.]; Lacon variabilis Cand. Australia
[Gahan] (BMNH).

I have not been able to trace any specimens determined as variabilis var. a or
var. b. by Candéze.

This species, which occurs in Queensland and South Australia, is the only represen-
tative of the subfamily recorded from New Zealand. Sharp (1877: 479), who
first reported the species from Auckland, suspected that it was imported by marine
traffic. The following records suggest that the species is now fairly well established.

New ZEALAND: N. Island. Auckland, 8 ex.; Papakura, 1I5.vili.24 and 30.v.39
(Richardson), 2 ex; Matakana, I1.iv.36 (Clarke), 1 ex; Wayby Gorge, 29.xii.1926
(Clarke), 1 ex; Ohakuna, 20.xi.1919 (Harris), 1 g, 1 2; Wellington, 2 ex; S. Island,
Waiwhero (Motueka), Nr. Neslon, 2.ix.1949, 1 J; Picton (Walker), 2 ex; Greymouth
(Helms). I 3; Blackball, x1i.1916 (Harris), 1 2 (BMNH).

Candéze (1891 : 24) places alternans MacLeay, 1872 in synonymy with variabilis
Candéze. The species is unknown to me so that the synonymy has not been con-
firmed. Neboiss (1956: 13), following Schenkling (1925: 30), lists maculosus
MacLeay, 1888 as a synonym of variabilis. This synonymy does not appear to
have been published previously and comparison of the descriptions suggests that it
is not justified. L. maculosus is smaller in size (about 7-4 mm) than variabilis
(10-12 mm) and possesses patches of reddish yellow scales and propleural tarsal
grooves, characteristics which do not occur in variabilis. I believe the catalogue
synonymy is due to a printer’s error, the line ‘maculosus M’Leay (nec Cand.)’, etc.,
being placed above the line ‘variatus nom. nov.’ instead of in its correct place below
it. Schenkling (1925 : 30) proposed L. variatus as a new name for L. variegatus
Schwarz, 1906, which is a junior secondary homonym of L. variegatus (Motschulsky,
1854). L. variegatus Schwarz was proposed as a new name for L. maculosus Mac-
Leay, 1888, a junior primary homonym of L. maculosus Candéze, 1874. Lacon
variatus Schenkling, 1925 is a junior primary homonym of L. variatus Candéze, 1890.
If maculosus MacLeay (type-material in MacLeay Museum, University of Sidney
according to Neboiss) is found to be a good species, it will require another new name.

Agrypnus variatus (Candéze) comb. n.

} Lacon variatus Candéze, 1890 : CXLVIII.
Adelocera variatus (Candéze) Fleutiaux, 1926 : 98.

228 C. M. F. von HAYEK

The description is based on a number of specimens collected by Cardon at Chota
Nagpore (see p. 273).

The type-material has not been located.

The generic attribution is based on the following specimen. Inp1A: 9, Chota
Nagpor. Bengal; L. variatus Cand. C. R. Soc. Ent. Belge. 1890: 148 [? Fleut.]
(MNHN, Paris). The specimen may be part of the syntype-series.

Agrypnus variolus (Candéze) comb. n.

Lacon variolus Candéze, 1874 : 92.
Lacon variolus (Candéze) Neboiss, 1956 : 14.

Lectotype (Van Zwaluwenburg, 1959 : 354, see p. 9). AUSTRALIA: 9, N. Holl.
Q’land; Janson coll. ex Candéze, 1903.130; Lacon variolus Cdz. n.sp. Queensland
[Cand.]; Lacon variolus Cand. Type ex coll. Cand. [Gahan] (BMNH).

The description is based on a number of specimens in the Candéze and Janson

collections. There are no more specimens from Janson collection in the BMNH.
The IRSNB, Brussels may possess additional specimens.

Agrypnus versicolor (Fleutiaux) comb. n.
Adeloceva (Archontas) versicolor Fleutiaux, 1932a : 51.

Holotype. 9, Mapacascar: Brickville; Cap Refroigney Colonel Grimlet; versicolor
Fleut. type [Fleut.] (MNHN, Paris).

Agrypnus vestitus Klug comb. n.

Elater (Agrypnus) vestitus Klug, 1833 : 66 (1834 : 154).

Lacon vestitus (Klug) Germar, 1840 : 264.

Lacon tumidicollis Candéze, 1857 : 108. [Synonymized by Candéze, 1874 : 53.]
Adelocera vestitus (Klug) Fleutiaux, 1926 : 96.

Elater (Agrypnus) vestitus Klug. LECTOTYPE (present designation). Mapa-
GASCAR: 4g, 16095 [Museum catalogue number = Madagascar, Goudot]; Madag.
Goud. (ZMHU, Berlin).

Paralectotypes. 3 9, Madagascar, Goud. Nr. 16095 (ZMHU, Berlin).

The specimens stand beside a blue Gerstaecker label: vestitus Germ. Elater
vestitus Klug. Madag. Goud. The absence of Klug’s label is probably due to
Gerstaecker (see p. 275).

Lacon tumidicollis Candéze. LECTOTYPE (present designation). 9, Laf.
Madagascar; Janson coll.; Lacon tumidicollis Cdze., Cand. type e coll. de Laferté;
Monstrosité individu de vestitus [Cand.} (BMNH).

GERMAR’S INTERPRETATION OF Lacon vestitus (KLUG). It is impossible to tell
from Germar’s work whether he owned specimens of this species or whether his

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 229

knowledge of the species was based on Klug’s description. Germar’s collection
was broken up (see p. 275). Neither the DEI, Eberswalde nor the NMHU, Berlin
contain specimens of vestitus Klug, petittt Dej. from ‘Cap’. The BMNH collection
contains a male and female from the Schaum collection. The male, which lacks
the abdomen, bears a small blue triangular label with the word ‘Mad.’ and a larger
blue label with the words ‘vestitus Kl., petitti Dej. Madag.’ in a handwriting I
believe to be that of Germar. The label also bears the words ‘Afr. mer.’ which
have been crossed out. The female bears a small blue triangular label with the
letters ‘Dp.’ probably standing for Dupont, and Candéze’s label ‘L. vestitus’. The
specimens are conspecific with the lectotype. The male was used in the preparation
of the table on p. 20. A. vestitus (Klug) has not been recorded from South Africa.

Agrypnus vicinus (Fleutiaux) comb. n.
Lacon vicinus Fleutiaux, 1919 : I9.

Adelocera vicinus (Fleutiaux) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). E. Arrica: g, Zanguebar (see p. 44);
265; Lacon vicinus Fleut., type [Fleut.]: Fleut. Voy. All. Jeann. Afr. or Elat. p.
[Fleut.] (MNHN, Paris).

Paralectotype. 4, Mandura, Zanguebar; Lacon vicinus Fleut., co-type [Fleut.]
(MNHN, Paris).

Agrypnus victoriae (Candéze) comb. n.

Lacon victoriae Candéze, 1865 : 12.
Lacon victoriae Candéze; Neboiss, 1956 : 14.

LECTOTYPE (present designation). AUSTRALIA: 9, N. Holl. Melbourne C.
Cdze; Janson coll. ex Candéze, 1903.130 Lacon victoriae Cdz. Austr. [Cand.]; Lacon
victoriae Cdze. Type ex coll. Cand. [Janson] (BMNH).

Paralectotype. 3, N. Holl. Melbourn. C. Cdze.; Janson coll. ex Candéze 1903.130
(BMNH).

Agrypnus viettei (Girard) comb. n.
Archontas viettei Girard, 1970 : 25.
Holotype. Mapacascar: g, massif du Tsaratanana, matsabory en dessous
de Andohanisambirano, 1,900 m [2,050 m], xii. 1964 (P. Soga). (MNHN, Paris).

Paratype. I g, same locality as holotype (Girard collection, Laboratoire de
Zoologie, Ecole normale supérieure, Paris).
Both specimens bear Girard’s determination labels.

| Lacon vitalisi Fleutiaux, 1918d : 190.
Adelocera vitalisi (Fleutiaux) Fleutiaux, 1926 : 96.

|

| Agrypnus vitalisi (Fleutiaux) comb. n.

230 Cc. M. F. von HAYEK

LECTOTYPE (present designation). NortTH VIETNAM: 3g, Region de Hanoi;
Lacon vitalisi Fleut., type [Fleut.] (MNHN, Paris).

Agrypnus wallacei (Candéze) comb. n.

Lacon wallacei Candéze, 1874 : 80.
Adelocevra wallace: (Candéze) Fleutiaux, 1926 : 96.

LECTOTYPE (present designation). SuLAWEsI: 3g, Tandano, Celebes; Janson
coll. 1903.130; L. wallacei Cdz. [Cand.]; Lacon wallacei Cand., type [Gahan] (BMNH).

Candéze (loc.cit.) recorded a variety from Macassar. The BMNH collection
contains 3 specimens from Macassar, Janson coll. One bears Gahans label ‘Lacon
wallacei Cand. Var.’. These specimens are not conspecific with the lectotype.
They belong to a species unknown to me.

Agrypnus yuppe (Kishii) comb. n.
Colaulon (Sagojyo) yuppe Kishii, 1964 : 31; pl. 2, figs 1, 2, 7, 9.
Sagojyo yuppe (Kishii) Ohira, 19676 : 103.
Holotype. JAPAN: Q, Is. Kuchi-us-erabu in Kagoshima Prefecture, July 29,
1963. Presumably in Kishii collection.

The generic attribution is based on the following specimen: 1 9, Hatsuno Amami-
Oshima, 28-VII. 1862; determined by Ohira, 1968 (BMNH).

Agrypnus zanzibaricus nom. n.

Tilotarsus [sic] marmoratus Candéze, 1878b : LIV (8).
Agrypnus marmoratus (Candéze) comb. n. [Junior secondary homonym of Agrypnus
marmoratus (Candéze, 1874).]

The description is based on material collected in Zanzibar in the course of Raffray’s
expedition.

Type-material: MCZN, Genoa according to Candéze (18780 : LIV).

The generic attribution is based on the following specimens. TANZANIA: I 4,
Zanzibar Raffray; Tilotarsus marmoratus Cdz. [Cand.]. 1 9, Zanzib.; Museum

Paris, Coll. Leon Fairmaire, 1906; Tilotarsus marmoratus Cdz. [Cand.] Cand. det.
(MNHN, Paris).

Agrypnus zietzi (Blackburn) comb. n.

Lacon zetzt Blackburn, 1985 : 48.
Lacon zietzi Blackburn; Neboiss, 1956 : 14.

The description is based on an unrecorded number of specimens taken at Lake
Callabonna (S. AUSTRALIA) by Mr Zietz. Unexpectedly (see p. 270) the species is
not represented in the BMNH collection.

According to Neboiss, the type-material is in the SAM, Adelaide.
The generic attribution is based on the description.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 231

MERISTHUS Candéze

Subgenus Meristhus Candéze, 1857 : 162. Type-species: Elater lepidotus Palisot de Beauvois,
by original designation.
Rhaciaspis Arnett, 1952 : 121. Type-species: Elater lepidotus Palisot de Beauvois, by original
designation.
Subgenus Sulcimerus Fleutiaux, 1947:255. Type-species: Meristhus quadripunctatus Candéze,
by subsequent designation (Arnett, 1955 : 617).

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae near the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepimeron and mesepisternum do not form part of
margin of mesocoxal cavity (Text-fig. 2). Second and third antennal segments cylindrical,
not triangular like the fourth and following segments (Text-fig. 13). Antennal groove extending
beyond the anterior half of the prosternopleural suture and deep enough to accommodate the
rolled antennae. Body clothed with scales. Prothorax not restricted behind the anterior
angles; lateral carinae attain the anterior margins. Propleurae with (Meristhus s. str.) or
without (subgenus Sulcimerus) depressions for the reception of the anterior tarsus. Scutellum
with distinct median longitudinal carina. Tarsi simple, without ventral lobes.

HISTORY OF THE GENUS. Candéze established the genus for those species in
which the scutellum bears a longitudinal carina.

There has been some difference of opinion concerning the type-species of Meristhus.
Golbach (1969a) discusses the problem, which probably stems from the fact that
not only did Hyslop (1921 : 656) designate scrobinula [sic] Candéze as the type-
species, but Candéze, who was not in the habit of designating type-species for the
genera he established, has already on two occasions made statements which, in my
opinion, amount to the designation of the type-species of Meristhus.

Candéze (1857 : 162), writing of Meristhus, states ‘ce petit genre, qui a pour
type l’espéce suivante, se distingue des Lacon et tous les autres Agrypnides par la
conformation tout particulaire de l’ecusson’. The first species, both in the key and
in the text is lepidotus Palisot de Beauvois. I believe that Fleutiaux (194Ie : 47)
was fully justified in accepting this statement as a valid designation of the type-
species of Meristhus. Forty years later Candéze (1897: 12), in describing M.
erinaceus from Borneo, remarks that this species is ‘un peu plus grande que la
M. scobinula [from China] type du genre...’. It seems probable that on this
occasion Candéze referred to scobinula as the type-species either because he has
forgotten his earlier statement or, because he was at that time dealing only with
oriental species. Hyslop’s designation is clearly not based on Candéze (1897) as
he states that it is a ‘present designation’. Why he selected this species is unknown,
but his choice may well have been influenced by the fact that scobinula occurs in
Mexico.

In the absence of intermediate forms, the retention of the subgenus Sulcimerus,
which includes those species possessing grooves on the propleurae for the accom-
modation of the anterior tarsi, appears justified at the present time. However,
when the four species at present unknown to me (see below) and additional material
belonging to the genus is available, it may become apparent that, as in Agrypnus,
this characteristic is of no value above the specific level.

SEXUAL DIMORPHISM. The sexes appear to be indistinguishable externally.

232 Gc. M. F. von HAYEK

DISTRIBUTION. Africa, Madagascar, India, Indonesia, Southern U.S.A.

BIOLOGY AND HABITS. The larvae of Meristhus (Sulcimerus) niponensis Lewis
live in the ground (Ohira, 1962). A Meristhus sp. near lepidotus Palisot de Beauvois
commonly occurs on the fore dunes of the eastern South African seaboard. The
adults have been found burrowing in the sand, usually in the neighbourhood of the
dune plants, on which the larvae (which are unknown) probably feed (Callan,
1964 : 128).

THE SPECIES INCLUDED IN THE GENUS

The Schenkling catalogue (1925 : 35) lists the names of 22 species, of which
two are regarded as synonyms of scobinula Candéze. Two species have been
described since the publication of the catalogue. The assignment of eight species
to Meristhus (s.1.) has been confirmed and the scobinula Candéze-texanus Horn
synonymy provisionally accepted. Fleutiaux (1947 : 457) transferred three species
to Rismethus and eight additional species are transferred to that genus in the present
work. The remaining four species, evinaceus Candéze, indecorus Candéze, ornatulus
Candéze and longicollis Candéze are unknown to me.

Meristhus (Meristhus) angulicollis Fairmaire
Meristhus angulicollis Fairmaire, 1897 : 382.

The description is based on an unrecorded number of specimens collected at
Suberbieville [MADAGASCAR] by Perrier.

Type-material: ? MNHN, Paris, or possible IRSNB, Brussels (see p. 275). Up to
the present time no specimens from the type-locality with Fairmaire’s determination
label have been located in the MNHN, Paris.

The generic attribution is based on the following specimens which may be syntypes;
MADAGASCAR: I 9, Suberbieville (Perrier); Meristhus angulicollis Fairm. t. Fleut.
I g, 1 2, Madg™, Suberb", H. Perrier; Museum Paris, Madagascar, coll. Perrier
de la Bathie. The female bears Fleutiaux’s determination label (MNHN, Paris).
2 ex, Suberbieville, 1903.28; Meristhus angulicollis Fairm. (BMNH). The registra-
tion number refers to a collection, including ‘paratypes’, purchased from Donckier.

Meristhus (Meristhus) lepidotus (Palisot de Beauvois)

Elatey lepidotus Palisot de Beauvois, 1805 : 11; Coleoptera, pl. 7, fig. 6.

Lacon lepidopterus Chevrolat, 1852 : 631. [Unjustified emendation.]

Meristhus lepidotus (Palisot de Beauvois) Candéze, 1857 : 163.

Meristhus lepidotus var. a (Agrypnus bohemanni Dej. in la Ferté Mus.) Candéze, 1857 : 163.

LECTOTYPE (present designation). NIGERIA: g, Benin; Janson coll. 1903.130;
Elater lepidotus P. de B., Agrypnus lepidotus Dej. ex coll. Dejean (Palisot de
Beauvois) [ Janson]; Meristhus lepidotus Pal. de Beauv. Cdze, type [ Janson] (BMNH).

Paralectotype: g, Benin, Meristhus lepidotus P. de B. Cdze, ex coll. Dejean
(Palisot de Beauvois) [Janson] (BMNH).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 233

According to Horn & Kahle (1925 : 201) Palisot de Beauvois’ collection passed
to the BMNH via Dejean, Janson and Godman and Salvin. The absence of de
Beauvois’ labels is probably due to Janson (see p. 276).

Fleutiaux (1919 : 15) states that de Beauvois’ type from Ovare [sic] cited by
Chevrolat (1852 : 631) is in his collection (now in the MNHN, Paris). In my
opinion this specimen (see below) cannot be the type. Palisot de Beauvois recorded
lepidotus from ‘prés de la ville de Benin’. He obviously regarded Oware as an
entirely different locality describing it as ‘pays voisin de la ligne et limitrophe du
Royaume de Benin’ (1805, introduction to the Flore de Oware et de Benin en Afrique).
Oware is almost certainly the district of Owerri in the Eastern Province of Nigeria.
The only specimen () in the Fleutiaux collection from the Chevrolat collection
and obviously the one to which Fleutiaux refers bears the following labels: 2 grey
labels with illegible numbers [? reference to locality list now lost]; lepidotus; Agryp-
nus lepidotus ? P.B. 11, pl. t.6 [next words illegible] ex Mus. Ol. [Chevrolat, according
to note by Fleutiaux on the underside of the label]; lepidotus Coll. Beauv. (coll.
Oliv.) Cand det., typique Mon. [Fleut.]; Fleut. Voy. Alluaud et Jeannel Elat. p.
[Fleut.]. Fleutiaux was probably misled into believing this to be de Beauvois’
type by Chevrolat’s statement (1852 : 630) that he had purchased part of Palisot
de Beauvois’ collection and the fact that the species was marked with an asterisk
in Chevrolat’s work indicating that it was represented in his collection. The speci-
men is conspecific with the lectotype.

Meristhus lepidotus var. a Candéze. LECTOTYPE (present designation).
SUDAN: Nubia; Janson coll. 1903.130; Meristhus lepidotus Pal. de B. var. Cdze,
Agrypnus bohemanni Dej. ex coll Dejean [Janson] (BMNH). The absence of
Candéze’s, la Ferté Sénectére’s or Dejean’s labels is probably due to Janson (see
p. 276).

Meristhus (Meristhus) squameus Candéze
Meristhus squameus Candéze, 1893 : 10.
The description is based on an unrecorded number of specimens collected at
Boma, Bas Congo [ZarRrE], by Tschoffen.
Type-material: ? IRSNB, Brussels.
The generic attribution is based on the description.

Meristhus (Sulcimerus) cristatus Horn comb. rev.

Meristhus cristatus Horn, 1871 : 299; pl. 4, fig. 2.
Rismethus cristatus (Horn) Golbach, 1969a :141.

Holotype. 4, dark steel-blue paper disk (? indicates California); Type, 8144;
Meristhus cristatus Horn [Horn] (MCZ, Harvard).

The description is based on a single specimen collected by John Xantus at Cape
San Lucas (between San José del Cabo and La Paz according to Horn (1894 : 302)),
California.

234 GM. Fe von HAYEK

For the explanation why this specimen is in the MCZ, Harvard and not the
ANS, Philadelphia, see note on the Xantus collection, p. 281.

Meristhus (Sulcimerus) insignitus Candéze
Meristhus insignitus Candéze, 1874 : 103.

LECTOTYPE (present designation): CEyLon: 9, Ceylon; Meristhus insignitus
Cdz. [Cand.]; Meristhus insignitus Cand. type [Janson]; sides of prothorax with
oblique fovae for reception of anterior tarsi [Janson]; Janson coll. 1903.130 (BMNH).

Paralectotype: 2, labels as lectotype but without Candéze’s determination label.

Meristhus (Sulcimerus) niponensis Lewis
Meristhus niponensis Lewis, 1894 : 30.

LECTOTYPE (present designation). JAPAN: 9, 6.9.81 [underside of card mount];
Niigata, 4.ix-16.ix; Meristhus niponensis Lewis, type [Lewis]; Japan, G. Lewis.
1910 : 320 (BMNH).

Lewis states that he had ‘several specimens’. Only the one designated as the
lectotype has been located. The others may have been given away by Lewis.

Meristhus (Sulcimerus) quadripunctatus Candéze
Meristhus quadvipunctatus Candéze, 1857 : 163.

LECTOTYPE (present designation). g, India bor. [Janson]; Janson coll.
1903.130; Meristhus quadripunctatus Cdze. Type ex coll. Laferté [ Janson] (BMNH).

The description is based on an unrecorded number of specimens from ‘Sylhet
[E. PAKISTAN] et Assam’ received from de la Ferté Sénectére and Deyrolle. The
discrepancy in the published locality and the label and the absence of Candéze’s
determination label is probably due to Janson (see p. 276). The material from the
Deyrolle collection has not been located.

Meristhus (Sulcimerus) scabrosus Fleutiaux

Meristhus scabrosus Fleutiaux, 1931 : 74.
Meristhus (Sulcimerus) scabrosus Fleutiaux, 1947 : 156.

LECTOTYPE (present designation). SourH VIETNAM: 9, Museum Paris,
Nhatrang, A. Krempf, 1913; Types; Meristhus scabrosus Fleut., types [Fleut.]
(MNHN, Paris). Length 6 mm.

Paralectotype, 2, length 6-5 mm, on the same card mount as the lectotype.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 235

RISMETAHUS Fileutiaux

Rismethus Fleutiaux, 1947: 257. Type-species: Meristhus scobinula Candéze, by original
designation.

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae near the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepisternum and mesepimeron do not form part of the
margin of the mesocoxal cavity (Text-fig. 2). Second and third antennal segments cylindrical,
not triangular like the fourth and following segments (Text-fig. 13). (Golbach, 1969a : 141,
corrected the printer’s error in Fleutiaux’s paper concerning the length of the second and third
antennal segments.) Antennal groove not extending beyond the anterior half of the prosterno-
pleural suture but deep enough to accommodate the rolled antennae. Body clothed with
scales. Prothorax constricted immediately behind the anterior angles; lateral carina does not
attain the anterior margin. Propleurae and metasternum with depressions for the reception
of the anterior and middle tarsi, but never with well defined grooves. Scutellum simple or
with a longitudinal carina. Tarsi simple, without ventral lobes.

The constriction of the prothorax immediately behind the anterior angles is the
only characteristic distinguishing the members of this genus from Agrypnus and
Meristhus. Another characteristic feature, the abbreviated lateral carina of the
prothorax also occurs in certain Agrypnus species, e.g. flavescens Candéze. As
more material becomes available it may become apparent that the shape of the
prothorax is not of sufficient importance to warrant the retention of the genus.
However, since no intermediate forms are known at the present time the genus
has been retained in this work.

HISTORY OF THE GENUS. The genus was established for species congeneric
with scobinula Candéze, which Fleutiaux considered could not be included in the
same genus (Meristhus) as lepidotus Palisot de Beauvois and quadripunctatus Can-
déze. According to Fleutiaux, the distinctive features of the genus are that the
scutellum is simple and that the metasternal tarsal depressions are less strongly
impressed than those on the propleurae. Study of the available material has
shown that these characters are of no importance at the generic level and the
genus is here redefined.

SEXUAL DIMORPHISM. The sexes appear to be indistinguishable externally.

DISTRIBUTION. Madagascar, Ceylon, ?Burma (‘Ind. or.’), Indonesia, China,
Samoa, Texas and Central America.

BIOLOGY AND HABITS. Nothing is known of the life history and habits of the
species included in the genus.

THE SPECIES INCLUDED IN THE GENUS RISMETHUS

Fleutiaux included three species in the genus. Golbach (1969a@ : 141) transferred
cristatus Horn and squanuiger Champion to the genus. The generic assignment
of the latter is confirmed. In my opinion there is no justification for the inclusion
of cristatus, in which the prothorax is not constricted behind the anterior angles,
in Rismethus. Seven species are newly assigned to Rismethus.

236 C. M. F. von HAYEK

Rismethus apicalis (Candéze) comb. n.
Meristhus apicalis Candéze, 1874 : 103.
LECTOTYPE (present designation). CELEBES: 9, Celebes/C.W.; Janson coll.

1903.130; ex coll. Wallace; Meristhus apicalis Cdz. [Cand.]; Meristhus apicalis
Cdze, type [Janson] (BMNH).

Paralectotype. 1 ex., Timor, C.W.; Janson coll. 1903. 130; Meristhus apicalis
Cdze. Cand. [Janson] (BMNH).

Rismethus diodesmoides (Motschulsky)

Brachylacon diodesmoides Motschulsky, 1861b : 118.
Meristhus biguttatus Candéze, 1893 : 10. [Synonymized by Fleutiaux, 1932e : 79.]
Rismethus diodesmoides (Motschulsky) Fleutiaux, 1947 : 257.

Brachylacon diodesmoides Motschulsky. LECTOTYPE (present designation)
, ‘C’ on yellow paper circle [? = CEYLON]; Brachylacon diodesmoides Motsch.
Ind. or. Ceylon [Motsch.]; Genus Brachylacon Cand., Fleutiaux det. [Fleut.] (ZMU,
Moscow).

Paralectotype: 2, yellow paper circle (ZMU, Moscow).
The description is based on an unrecorded number of specimens from Ceylon.

Mensthus biguttatus Candéze. The description is based on an unrecorded number
of specimens from Perak [? Island or region W. Mataysia].

Type-material: IRSNB, Brussels, according to Fleutiaux (1947 : 237).

The acceptance of the synonymy is based on the examination of a number of
specimens from Perak standing as biguttatus in the BMNH. These specimens
may be part of the syntype series.

Rismethus erinaceus (Candéze) comb. n.

Lacon evinaceus Candéze, 1874 : 74.
Meristhus evinaceus (Candéze) Fleutiaux, 1926 : 97.

LECTOTYPE (present designation). BorNEO: g, Sarawak/C.W.; Janson coll.
1903 : 130; Lacon erinaceus (type) Cand. [Cand.]; Lacon erinaceus Cand., type,
Sarawak [Gahan] (BMNH).

Rismethus minusculus (Candéze) comb. n.
Meristhus minusculus Candéze, 1874 : 104.

LECTOTYPE (present designation). BorNEo: Sarawak, bearing Candéze’s deter-
mination label (MCSN, Genoa).

Not examined, details of labels unknown to me.

The generic attribution is based on a report on the specimen kindly supplied
by Dr D. Guigha.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 237

Rismethus mocquerysi (Fleutiaux) comb. n.

Meristhus mosquerysi Fleutiaux, 1899 : 223.

LECTOTYPE (present designation). MADAGASCAR: 9, Baie d’Antongil; Meristhus
mocquerysi Fleut., type [Fleut.] (MNHN, Paris).

Rismethus nigritulus (Candéze) comb. n.
Meristhus nigritulus Candéze, 18930 : Io.
Lectotype (van Zwaluwenburg, 1959 : 355, see p. 9). SUMATRA: sex and
details of labels not recorded. Van Zwaluwenburg’s comment ‘Candéze’s designa-

tion’ suggests that the specimen bears Candéze’s determination label with the
word ‘type’. (RSNB, Brussels). Not examined.

Paralectotypes: 2 ex., recorded by Van Zwaluwenburg (IRSNB, Brussels). Not
examined.

The generic attribution is based on the following specimens which may be part
of the syntype-series: I ex., Sumatra; Meristhus nigritulus Cand. Cand det [Cand.].
I ex., Palembang, Sumatra (MNHN, Paris).

The published locality is Sumatra, Palembank [sic].

Rismethus oceanicus (Van Zwaluwenburg) comb. n.
Meristhus oceanicus Van Zwaluwenburg, 1945 : 95; fig. I.
Holotype. Samoa: ? 9, government garden, Taputimu, Tutuila, Samoa, Aug.

7 at 100 ft, sweeping sweet potato plants. Zimmerman (BPBM, Honolulu). Not
examined.

Paratypes: 3 ex., locality as lectotype (BPBM, Honolulu) 1 ex, ibidem (HSPA,
Honolulu). Not examined.

The generic attribution is based on the description.

Rismethus perraudieri (Fleutiaux)

Meristhus perraudieri Fleutiaux, 1889 : 139.
Rismethus perraudieri (Fleutiaux) Fleutiaux, 1947 : 255.

LECTOTYPE (present designation). SouTH VIETNAM: 9, QN; Meristhus
perraudieri Fleut., type, Quinhon Annam [Fleut.] (MNHN, Paris).
The published locality is Qui-Nhon.

Rismethus pistrinarius (Candéze) comb. n.

Meristhus pistrinarius Candéze, 1857 : 164.

LECTOTYPE (present designation). 4, Ind. bor. [sic., Janson]; 49; Janson
coll. 1903.130; Meristhus pistrinarius Cdze, type ex. coll. Laferté [Janson] (BMNH).

238 CUM. EF. von SHAY EK

Paralectotype: I ex., Ind. bor. [sic., Janson]; 49; Janson coll. 1903.130; Meristhus
pistrinarius; ex coll. Laferté [both Janson] (BMNH).

The description is based on an unrecorded number of specimens from ‘Des Indes
Orientales’ from the La Ferté and Deyrolle collections. The discrepancy between
the published locality and the labels is almost certainly due to Janson (see p. 276).
The specimens from the Deyrolle collection have not been located.

Rismethus scobinula (Candéze)

Meristhus scobinula Candéze, 1857 : 164; pl. 2, fig. 26.
Meristhus texanus Horn, 1871 : 300; pl. 43, fig. 1. [Synonymized by Horn, 1875 : 148.]
Rismethus scobinula (Candéze) Fleutiaux, 1947 : 257.

Meristhus scobinula Candéze. Candéze states that this species is found in China
and Mexico but his comments in the last paragraph on page 165 concerning Chevro-
lat’s Mexican specimen suggest that the description is based on a specimen or
specimens from China. The remarks by Lewis (1894 : 30) ‘it appears clear that
his type example came from China’ adds support to this opinion. Fleutiaux’s
(1919 : 15) statement that he possesses the type of Candéze’s Monograph, a specimen
from Teapa (Mexico) sent by Pilate to Chevrolat cannot be accepted as a valid
lectotype designation.

LECTOTYPE (present designation). CHINA: g, China; Janson coll. 1903.130;
ex coll. Candéze; Meristhus scobinula Cdz., Chine [Cand.]; scobinula Cdze, type
Mon. [Janson] (BMNH).

Meristhus texanus Horn. LECTOTYPE (present designation). U.S.A.: 9,
Tex.; Lectotype 3830 [Museum curatorial label, designation unpublished] Meristhus
scobinula Cdz., texanus Horn [?Horn] (ANS, Philadelphia). The description is
based on two specimens from Texas collected by Belfrage. The second specimen
cannot be found in the ANS, Philadelphia. If it is not in Belfrage’s collection
(USNM, Washington according to Horn & Kahle, 1935: 15), I believe that the .
following specimen should be regarded as the Paralectotype: 2, Texas, 74.12 [BMNH |
registration number = Belfrage, Texas]; Meristhus texanus [Horn] (BMNH).

The comparison of the external characteristics and male genitalia of Chinese
and American specimens in the BMNH has served only to confirm the synonymy.
However, the distribution, which is of a type unusual in the Elateridae, suggests
that a critical investigation into the structure, life history and habits of individuals
from the two regions is desirable.

Candéze (1891 : 26) placed M. setarius Chevrolat, 1867 : 596 in synonomy with
scobinula Candéze. The description is based on an unrecorded number of specimens
from ‘Cuba. Collections de MM. Gundlach et Poéy’. According to Horn & Kahle
(1935 : 100) the Gundlach collection is in the Instituto de Ensenanza de la Habana,
Cuba (not examined) and the Poéy collection (Horn & Kahle, 1936: 211) passed
to the American Entomological Society, Philadelphia (not examined). Chevrolat
(1864 : 425) acquired Poéy’s first Cuban collection and also duplicates from later

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 239

collections but these specimens cannot be found in his collection in the MNHN,
Paris.

Lewis (1894 : 30) recorded scobinula from Japan. The BMNH collection contains
three specimens, one with Lewis’s determination label, from the Lewis collection.
These specimens, in which the metathorax is reduced in length and wings are
absent, are not conspecific with the lectotype of scobinula Candéze.

Rismethus squamiger (Champion)

Meristhus squamigey Champion, 1894 : 269; pl. 11, fig. 14.
Rismethus squamigey (Champion) Golbach, 1969a@ : 141.

LECTOTYPE (present designation). Right hand of two specimens on one
card, right antenna missing, GUATEMALA: 3, Duenas, Guatemala, G. C. Champion;
BCA coll. III(r) Meristhus [printed] squamiger [Champ.]; sp. figured (BMNH).

Paralectotypes: Specimen beside lectotype and 7 ex, with same locality and
determination labels as the lectotype (BMNH). 2 ex, ibidem (MNHN, Paris).
2 ex, Guatemala City, 500 ft, Champion; determination labels as lectotype. 2 ex
on one card, San Geronimo, Vera Paz, Champion; determination label as lectotype.
I ex, Capetillo, Guatemala, G. C. Champion; determination label as lectotype.
I ex, Santarem, Bates coll.; Janson coll. 1903.130; squamiger Ch. [Champ.]; Meristhus
scobinula Cdz. [Janson]. 2 ex, Teapa, Mexico; Janson coll. 1903 : 130; squamiger
Champ. [Champ.]; Meristhus scobinula Cdze [Janson] (BMNH).

TRIERS Candéze

Trievs Candéze, 1900 : 78 (3). Type-species: Tviers vamitarsas Candéze, by monotypy.

GENERIC DIAGNOSIS. Each tarsal claw with a group of setae near the base (Text-fig. 11).
Tibial spurs absent (Text-fig. 10). Mesepisternum and mesepimeron do not form part of the
margin of the mesocoxal cavity (Text-fig. 2). Second and third antennal segments small,
subequal, each smaller than the fourth and following segments (Text-fig. 13). Antennal groove
not extending beyond the anterior two-thirds of the prosternopleural suture. Body clothed with
scales. Prothorax not constricted behind the anterior angles; lateral margin of the prothorax
carinate. Propleurae and metasternum without distinct grooves or depressions for the
reception of the tarsi of the anterior and middle legs. Scutellum without longitudinal carina.
First four tarsal segments expanded laterally (Text-fig. 17).

HISTORY OF THE GENUS. The genus was established for a single species from
Australia which differs from all other known Agrypninae in that the first four
tarsal segments are expanded laterally.

SEXUAL DIMORPHISM, BIOLOGY AND HABITS. Nothing is known of the life history
and habits of this species. It is not known whether this species displays sexual
dimorphism.

THE SPECIES INCLUDED IN THE GENUS

| At the present time only one species is known.

240 CG. M: EF. von HAYEK

Triers ramitarsus Candéze
Trievs vamitarsus Candéze, 1900 : 78 (3).

The description is based on an unrecorded number of specimens from Australia,
probably from the northern part of the continent.

Type-material: IRSNB, Brussels according to Neboiss (1956 : 15). Not confirmed.

The generic attribution is based on the following specimen; AUSTRALIA: 4,
Northern Territory; BM 1968.196; determined from the description, CMFH (BMNH).

LANELATER Arnett

[Agrypnus sensu auct., nec Eschscholtz, 1829. Misinterpretation.]

Amaurus Castelnau, 1840 : 23. Type-species: Amaurus senegalensis Castelnau (=Lanelater
notodonta (Latreille)), by subsequent designation (Hyslop, 1921 : 625). [Homonym of
Amaurus Burmeister, 1835, Hemiptera. ]

Lanelatey Arnett, 1952 : 105. Type-species: Agrypnus schotti LeConte, by original designation.

GENERIC DIAGNOSIS. Each claw bearing a group of setae near the base (Text-fig. 11).
Tibial spurs present (Text-fig. 9). Mesepisternum and mesepimeron form part of the margin
of the mesocoxal cavity (Text-fig. 1). The third antennal segment never resembling the fourth
in shape, but often considerably longer than the second segment. Antennal groove variable
in depth but extending at least three-quarters of the length of the prosternopleural suture.

Body clothed with setae; scales are entirely absent. Propleurae and metasternum without
depressions or grooves for the accommodation of the anterior and middle tarsi. Prothorax
simple, without a constriction behind the anterior angles; lateral carinae present. Scutellum
simple, without longitudinal carina. Tarsi simple, without ventral lobes.

RANGE OF VARIATION FOUND WITHIN THE GENUS. In general the species included
in the genus bear a close resemblance to one another. The greatest range of varia-
tion is found in the antennal groove, which varies considerably in length and depth
from one species to another.

HIsTORY OF THE GENUS. Arnett established the genus Lanelater for ‘those
species which were formerly placed in the genus Agrypnus’ sensu auct., nec
Eschscholtz, 1829 (type-species Elater tomentosus Fabricius, see below) but which,
since Lane’s (Arnett, 1952 : 105) discovery that Westwood (1838, Synopsis, p. 26)
had properly designated Elater murinus Linnaeus as the type-species of Agrypnus
(see p. 114), could no longer be accommodated in that genus. The next available
generic name for these species is Amaurus Castelnau, 1840 (type-species Amaurus
senegalensis Castelnau) but this name is pre-occupied by Amaurus Burmeister,
1835.

The interpretation of Agrypnus as a genus containing only species congeneric
with tomentosus can be traced to Candéze (1857, see p. 114). Before that time
the genus was poorly defined and in addition to murinus and tomentosus, included
species now assigned to Lacon (e.g. atomarius Fabricius = punctatus Herbst),
Danosoma (fasciatus Linnaeus), Alaus (moerens Germar) and Aliteus (adspersus
Herbst). Hyslop’s designation of tomentosus as the type-species was probably
made not so much because it heads Eschscholtz’s list of species but because it
corresponded with the interpretation of Agrypnus at that time.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 241

SEXUAL DIMORPHISM. In the majority of species there are no obvious differences
between the sexes. In a few species (e.g. aequalis (Candéze)) the antennae of the
males are longer than those of the females. In some other species (e.g. babaulti
(Fleutiaux)) the females are larger and have a stouter appearance than the males.

DISTRIBUTION. The genus is represented in all parts of the world except South
America.

BIOLOGY AND HABITS. The early stages of the larva of L. fuscipes (Fabricius)
have been described and figured by Fletcher (1919). The larvae of this species
and of L. mastersi (Macleay) are predaceous and may be of some economic impor-
tance. It seems probable that the larvae of fuscipes attack larvae of Oryctes
rhinoceros Lesne in Mauritius and the larvae of mastersi are believed to play an
important part in keeping down the numbers of Rutelid and Melolonthid larvae,
which are serious pests of sugar-cane plantations in Australia (Illingworth, 1g2r : 16).

THE SPECIES INCLUDED IN THE GENUS

Arnett states that the genus is moderately large and that current catalogues
list 80 species (Palaearctic, 9; Indomalaysian, 30; Australian and Oceanic, 7; Africa
and Madagascar, 31; Americas, 3). It is not immediately clear how Arnett arrives
at these figures. The most probable explanation appears to be that they are based
on the Schenkling (1925) catalogue of Agrypnus without taking into account the
species which appear more than once (e.g. aegualis Candéze occurs both in the
Indomalaysian region and Australia) or the corrections and additions listed in the
addenda and corrigenda at the end of the second volume. Schenkling lists the

following number of species in each section: Palaearctic, 9; Indomalaysian, 28;
Australian and Oceanic, 7; Africa and Madagascar, 28 and the Americas, 3.
With the addition of two Indomalaysian species (lopezei Fleutiaux, 1934 and
pescadoriensis Miwa, 1934) and three African species (accwminatus, confusus
| and proximus Fleutiaux, 1935) described subsequently, these figures agree with
| Arnett’s.
| In the catalogue which follows, the species listed in the uncorrected Schenkling
| catalogue together with the five above mentioned species are assumed to have
_ been transferred to Lanelater by Arnett, 1952: 105. All others are recorded as
new combinations.

Lanelater is the only genus included in the Agrypninae in which scales are entirely
absent. This characteristic together with the long antennal grooves and the
presence of tibial spurs makes the generic attribution of specimens and in many
cases of species known only from the description a relatively simple matter. It
has therefore been considered unneccessary to record the data on the specimen

or specimens on which the generic attribution is based.

.
. | Lanelater acuminatus (Fleutiaux)

Agrypnus acuminatus Fleutiaux, 1935¢ : 90.
Lanelater acuminatus (Fleutiaux) Arnett, 1952 : 105.

Q

f

242 € M. F. von HAYEK

LECTOTYPE (present designation). East Arrica, location unknown: 4,
McArthur, Thua R., Nov. 33; Agrypnus acuminatus Fleut., type [Fleut.] (MNHN,
Paris).

The locality Thua River does not appear in any gazetteer known to me. The
BMNH collection contains specimens, believed to be conspecific with the lectotype,
from KrEnyA, Turkana.

Lanelater aequalis (Candéze)

Agrypnus aequalis Candéze, 1857 : 25.
Agrypnus insularis Fairmaire, 1891 : LXX. [Synonymized by Fleutiaux, 1923 : 409.]
Lanelater aequalis (Candéze) Arnett, 1952 : 105.

Agrypnus aequalis Candéze. LECTOTYPE (present designation): 9, C.M.
[probably indicating ‘Collection Mniszech’] type [? Cand.]; specimen typiq. sur
laquel a eté etablie ’espéce 1857 [Cand.]; Agrypnus aequalis Cand. det. E. Candéze
[IRSNB curatorial label] (IRSNB, Brussels).

The description is based on an unrecorded number of specimens from “Des Indes
Orientales’ in the Mniszech collection. The reason for absence of Candéze’s deter-
mination label is unknown.

Agrypnus insularis Fairmaire. LECTOTYPE (present designation). SEYCHELLES:
9, Seychel; Agrypnus insularis Fairm. Seychelles [Fairm.]; Type [? Fairm.]; C.
Fairm.; Agrypnus insularis Fairm. det L. Fairm.; Type, insularis; Sec S. Schenkling
Col. Cat. Junk XI. 1935-27, p. 4. Agrypnus aequalis Cand. [the last three labels
are all modern IRSNB curatorial labels} (IRSNB, Brussels).

Paralectotypes. I 4, insularis Fairm. Seychelles [Fairm.]; sondaicus var. loc.
insularis Frm. Seychelles [Cand., blue border]; insularis Fairm. Seychelles [Fairm.];
Agrypnus insularis Fairm. det. E. Fleutiaux [IRSNB curatorial label]; collection
E. Candeze. 1 4, Seychelles; C. Fairm.; Agrypnus insularis Fairm. det L. Fairmaire
[IRSNB curatorial label]; Collection E. Candeze (IRSNB, Brussels). 1 9, Seychel.;
Agrypnus insularis Seychelles [Fairm.]. Museum Paris, Collection L. Fairmaire,
1906; Type. 1 9, Seychelles, Fallou. Agrypnus insularis Fairm. Seychelles;
Museum Paris, coll. L. Fairmaire, 1906. 1 9, insularis Frm. Seychelles [Fairm.];
Museum Paris, coll. L. Fairmaire, 1906. 1 9, Mahe, Fallou; Museum Paris coll.
L. Fairmaire 1906. (MNHN, Paris).

The description is based on an unrecorded number of specimens collected in the
Seychelles [no detailed locality given] by R. P. Phillibert and submitted by M. G.
Fallou. Fleutiaux (1923: 409) states that the type is in the IRSNB, Brussels,
but as he provides no means of identifying the specimen his statement cannot be
accepted as a valid lectotype designation.

The MNHN, Paris contains a number of specimens with a printed locality label:
Iles Seychelles, La Digue, Ch. Alluaud, 1892. Several bear Fleutiaux’s determina-
tion labels; Agrypnus insularis Fairm. ex type. However if the specimens were
collected by Alluaud in 1892 they cannot have formed part of the original series.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 243

Lanelater andrewsi (Candéze)

Agrypnus andrewsi Candéze, 18934 : 169.
Lanelater andrewsi (Candéze) Arnett, 1952 : 105.

Holotype. Inp1A: 9, Belgaum S.; 997; Agrypnus andrewsi Cand. [Andrewes];
Agrypnus bipunctatus Cand; Andrewes Bequest 1922 : 221 (BMNH).

Andrewes appears to have replaced Candéze’s determination label with his own.
The synonymy with bipunctatus Candéze has not been published. It is probably
correct but more specimens of both sexes are required before the synonymy can
be confirmed.

Lanelater arabicus (Candéze)

Agrypnus arabicus Candéze, 1874 : 6.
Lanelater avabicus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). ARABIA: 3, Arabia; Janson coll. 1903.130;
arabicus type unique [Cand.]; Agrypnus arabicus Cdze., type ex Castelnau [ Janson]
(BMNH).

Candéze did not record the fact that he based the description on a single specimen.

Lanelater arizonae (Candéze)

Agrypnus arvizonae Candéze, 1897 : 5.
Lanelater avizonae (Candéze) Arnett, 1952 : 105.

Holotype. U.S.A.: Arizona, Tucson (Wickham) (MCSN, Genoa according to
Candéze, loc. cit. Arnett’s statement that the type is probably in the Geneva
Museum is almost certainly erroneous). Not examined.

The generic attribution is based on a series of specimens (10 3, I 9) from Tucson,
Arizona (Bryant) in the CAS, San Francisco determined by M. C. Lane. The
female agrees well with the holotype of schotti LeConte and I suspect that examina-
tion of the type of avizonae will show that the two species are conspecific.

The four (male) specimens (from Globe, Arizona) standing as arizonae in the
‘Fall collection (MCZ, Harvard) recorded by Arnett (1952: 107) are conspecific
with the specimens from Tucson. They differ from Arnett’s key in that the antennal
grooves are not uniformly deep but become more shallow towards the base.

Lanelater attenuatus (Candéze)

Agrypnus attenuatus Candéze, 1874 : 4.
Lanelater attenuatus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). West Arrica: g, Niger source [Janson];
Janson coll. 1903.130; attenuatus Cdz. [Cand.]; Agrypnus attenuatus Cdze, type
[Janson] (BMNH).

Paralectotypes. 1 9, Niger source [Janson]; Janson coll. 1903.130; attenuatus

244 CoM. FC von “HAYEK

Cdz. [Cand.]; Agrypnus attenuatus Cdze, Cand. [Janson]. 1 4, 1 9, same data
but without Candéze’s determination labels (BMNH).

The published locality is “Guinée [see Elasmosomus cornutus (Candéze) p. 103]
bouches du Niger.’ The discrepancy between the published locality and that on
the labels probably arose when Janson relabelled the specimens.

Lanelater attonitus (Candéze)

Agrypnus attonitus Candéze, 1874 : Io.
Lanelatey attonitus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). InpD1A: 9, India; Janson coll. 1903.130;
A. attonitus type [Cand.]; Agrypnus attonitus Cdze, type [Janson] (BMNH).

Candéze comments that though labelled ‘India’ he believes that the species
comes from the Malayan peninsula (‘presqu’ile Hindoue’). As there are no
additional records of the species this has not been confirmed.

Lanelater australis (Candéze)

Agrypnus australis Candéze, 1874 : 5.
Lanelater australis (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). Botswana: 92, Lake N’Gami; Janson
coll. 1903.130; A. australis Cdz., type L. N’Gami [Cand.]; Agrypnus australis
Cdze, type ex coll. Castelnau [Janson] (BMNH).

Lanelater babaulti (Fleutiaux)

Agrypnus babaulti Fleutiaux, 1918c : 168.
Lanelater babaulti (Fleutiaux) Arnett, 1952 : 105.

The original description is based on an unrecorded number of specimens, with
a size-range of 26-27 mm, collected by Babault in E. Africa. Three years later
Fleutiaux (1921: 5) published a slightly expanded description of babault1 and
recorded two specimens collected by Babault in the Kedong Valley.

LECTOTYPE (present designation). KrNnya: J, Kedong Valley (B.E.A.);
G. B. Mars 1913; Agrypnus babaulti Fleut., cotype, 1918 [Fleut.]; infuscatus KI.
comparé au type, Mus. Berlin [Fleut.] (MNHN, Paris).

Paralectotype. 1 9, Kedong Valley (B.E.A.); G. B. Mars, 1913; Agrypnus
babaulti Fleut., Type, 1918 [Fleut.]; Esp. fig. (MNHN, Paris). Aedeagus missing.

The specimen labelled as the type by Fleutiaux has not been selected as the
lectotype because the aedeagus is missing.

The Kedong Valley is about 35 miles east of Nairobi.

Fleutiaux did not publish the synonymy with infuscatus Klug. The two species
are not synonymous.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 245

Lanelater badeni (Candéze)

Agrypnus badeni Candéze, 1889 : 68 (2).
Lanelater badeni (Candéze) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens from CAMEROUN,
Isulu.

Type-material: ?>IRSNB, Brussels.
The confirmation of Arnett’s generic attribution is based on the description.

Lanelater bartoni (Fleutiaux)

Agrypnus bartoni Fleutiaux, 19024 : 163.
Lanelater bartoni (Fleutiaux) Arnett, 1952 : 105.

LECTOTYPE (present designation). ¢, InpIA: Agrypnus bartoni Fleut. type
[Fleut.] (MNHN, Paris).

Paralectotypes. 2 ex, India. 1 g, C. India, Ajmer, Rajputana. These
3 specimens stand beside the lectotype in the Fleutiaux collection but lack
determination labels. 1 3, T. R. Bell, Karachi; bartoni Fleut. [Fleut.] (MNHN,
Paris). 19, T. R. Bell, Karachi, Andrewes Bequest 1922 : 221; Agrypnus bartoni
Fleut. [Fleut.] (BMNH).

The published locality is ‘India’ without further details. T. R. D. Bell collected
in Kanara in the 1890s and sent the collection to H. E. Andrewes at the end of the
century (Kinnear, 1948). The material would therefore have been available to
Fleutiaux in 1902 and the inclusion of these specimens in the syntype-series appears
justified. There are probably a number of as yet unrecognized paralectotypes in
existence as none of those recorded above are 24 mm long, the maximum length
recorded by Fleutiaux.

Lanelater bifoveatus (Candéze)

Agrypnus bifoveatus Candéze, 1857 : 41.
Lanelater bifoveatus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). PuHitippInes: 3g, Dej. Manilla; Janson
coll. 1903 : 130; Agrypnus bifoveatus Cdze, Cand., type (A. tomentosus Dej. Cat.)
(e coll. Dejean) [Janson] (BMNH).

Paralectotype: 1 3, Dej. Manilla; Janson coll. 1903 : 130; Agrypnus bifoveatus
Cdze, Cand. v. (A. tomentosus Dj. Cat.) (e coll. Dejean) [Janson] (BMNH).

The specimen in the Deyrolle collection bearing this name, to which Candéze
refers, has not been found (see p. 274).

Lanelater bipunctatus (Candéze)

Agrypnus bipunctatus Candéze, 1857 : 29.
Lanelater bipunctatus (Candéze) Arnett, 1957 : 105.

246 C. M. F. von HAYEK

Holotype. 3 [aedeagus lost]; India bor., de Laf.; Janson coll. 1903 : 130;

A. bipunctatus [Cand.]; Agrypnus bipuntatus Cdze, Cand. type (e coll. de Laferté)
[Janson] (BMNH).

The published locality is Sylhet (E. PAKISTAN).

The antennae are malformed. In the left-hand antenna the 5 + 6, 7+ 8 and

9 + 10 segments are fused and in the right-hand antenna the 5 + 6 and 7+ 8
segments are fused.

Lanelater bradshawi (Candéze)
Agrypnus bradshawi Candéze, 1889 : 68 (2).
Lanelatey bradshawi (Candéze) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens from Zambezi
found by Bradshaw.

Type-material: ?IRSNB, Brussels.

Candéze records a specimen in the museum of the Societé Natura Artis Magistra.
This collection is now in the ZM, Amsterdam, but Dr J. P. Duffels tells me that
the specimen cannot be found in that collection.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater cinereus (Candéze)

Agrypnus cinereus Candéze, 1857 : 40.
Lanelater cinereus (Candéze) Arnett, 1952 : 105.

The description is based on a single headless specimen from ‘des Indes Orientales’
in the Deyrolle collection (see p. 271). The specimen has not been located.

The confirmation of Arnett’s generic diagnosis is based on material standing
as cinereus in the BMNH.

Lanelater confusus (Fleutiaux)

Agrypnus confusus Fleutiaux, 19350 : 195.
Lanelater confusus (Fleutiaux) Arnett, 1952 : 105.

LECTOTYPE (present designation). KENyA: 9, Kenya, Lokitang, Turkana
Nord, 750m; Agrypnus confusus Fleut., type [Fleut.] (MNHN, Paris). Length:
27 mm.

Paralectotype: 92, same locality as lectotype, labelled ‘paratype’ but without
a determination label (MNHN, Paris). Length: 25 mm.

Fleutiaux records a size-range of 22-27 mm. The smaller specimen or specimens
have not been located.

Lanelater crassiusculus (Candéze)

Agrypnus crassiusculus Candéze, 1857 : 30.
Lanelater crassiusculus (Candéze) Arnett, 1957 : 105.

|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 247

The description is based on a single specimen from SENEGAL submitted by
Mniszech.

Holotype. ?IRSNB, Brussels.
Confirmation of Arnett’s generic attribution is based on the description.

Lanelater crassiventris (Schwarz)

Agrypnus crassiventris Schwarz, 1899 : 75.
Lanelater crassiventris (Schwarz) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens with a size-
range of 19-21 mm from Deutsch Ost-Africa [TANZANIA] collected by Bennigsen.

Syntypes: ?>DEI, Eberswalde.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater dewalquii (Candéze)

Agrypnus dewalquit Candéze, 1857 : 34.
Lanelater dewalquii (Candéze) Arnett, 1957 : 105.

LECTOTYPE (present designation). ANcGoLa: g, Dey. Benguela; Janson
coll. 1903.130; A. dewalquii [Cand.]. Agrypnus dewalquii Cdze, Cand., type
(e coll. Deyrolle) [Janson] (BMNH).

Lanelater divergens (Fairmaire) comb. n.
Agrypnus divergens Fairmaire, 1892 : rot, 80.

Holotype. F.T.A.I. [previously French Somaliland]: ¢, divergens Fairm. Obock
[Fairm.]; Le Moult vend; Agrypnus divergens Fairm. [IRSNB curatorial label];
R. Mus. Hist. Nat. Belg 1.G.12.595 (IRSNB, Brussels).

The description is based on a single specimen, collected by Dr Gaujon between
3 April, 1890 and 7 April, 1891, and submitted by Maurice Aubert. The above,
the only specimen bearing Fairmaire’s determination known to me, was presumably
acquired by Le Moult when Aubert’s collection was broken up (Horn & Kahle,
hOQ25: 7).

This species is not listed in the main body of the Schenkling (1925) Catalogue
and is assumed to have been overlooked by Arnett (1957). See p. 241.

Lanelater ereptus (Candéze)

Agrypnus mastersi var. C., Agrypnus ereptus Jans., mss. Candéze, 1874 : 13.
Lanelatey eveptus (Candéze 1874) Van Zwaluwenburg, 1959 : 349.

Lectotype (designated by Van Zwaluwenburg, 1959 : 349). N.W. AUSTRALIA:
2, Nicol Bay/N.W. Australia; Janson coll. 1903.130; Agrypnus ereptus mihi

248 Cc. M. F. von HAYEK

>

[Janson]; Specimen communicated to Candéze and returned as ‘Agryp. Mastersii
Mc.L. var. C. A. ereptus Jans. Mss’ [Janson] (BMNH).

The name erepitus, which was first published in synonymy (see above), was made
available by Van Zwaluwenburg’s treatment of it as an available name with its
original authorship and date, and his adoption of it as the name of a taxon (IRZN,
Article 11(d), 1963 emendation).

Lanelater fallaciosus (Fairmaire) comb. n.

Agrypnus fallaciosus Fairmaire, 1892 : Iol.

LECTOTYPE (present designation). F.T.A.I. [previously French Somaliland]:
2, Obock (Aubert); fallaciosus Fairm. n.sp. [Fairm.] (MNHN, Paris).

Paralectotype: 1 9, Obock; fallaciosus Frm. type [Fairm, date illegible]; Museum
Paris. Coll. Abeille de Perrin (MNHN, Paris).

The species was described (loc. cit.: 80) as ‘assez commun’ but only the above
two specimens have been located.

This species is not listed in the main body of the Schenkling (1925) catalogue
and is assumed to have been overlooked by Arnett (1957). See p. 241.

Lanelater fleutiauxi nom. n.
Agrypnus eveptus Fleutiaux, 1902@: 165. [Junior primary homonym of Agrypnus ereptus

Candéze, 1874 : 13.]
Lanelatey eveptus (Fleutiaux) Arnett, 1952 : 105.

LECTOTYPE (present designation). AUSTRALIA: 9, Australie; Sud. Austr.;
A. ereptus Jans. in litt. [Cand.] Candéze det. [Fleut.]; A. ereptus Fleut., type
[Fleut.]; ereptus Fleut., Bull. Soc. Ent. Fr. 1902 p. 164 [Fleut.] (MNHN, Paris).

Van Zwaluwenburg (1959 : 349) comments that Fleutiaux’s description does
not agree with the lectotype of eveptuws Candéze. The explanation is that although
Fleutiaux refers to A. mastersi var. c Candéze = eveptus Jans. in litt.’ he based
his description not on Candéze’s specimen from Nicol Bay, N.W. Australia (see
p- 247) but on a specimen from an unknown locality in S. Australia in his own
collection. The two specimens are not conspecific.

Lanelater funestus (Candéze) comb. n.

Agrypnus funestus Candéze, 1857 : 35.
Agrypnus funestus Candéze; Fleutiaux, 1926 : 93. [Restored tospecificstatus. Notasynonym
of aequalis Candéze.]

The description is based on an unrecorded number of specimens from ‘Des Indes
Orientiales’ (see p. 271). There are no specimens from this locality with Candéze’s
determination label in the BMNH or IRSNB, Brussels.

The interpretation of the species is based on the following specimen, the only
one with Candéze’s determination label known to me: 1 3g, Deyr. India; Janson

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 249

coll. 1903.130; A. funestus [Cand.]; Agrypnus funestus Cdze, Cand. e coll. Deyrolle
[Janson] (BMNH). The specimen which may be a syntype, is not conspecific
with A. moestus Candéze and giluus Candéze as recorded by Fleutiaux (1926 : 93).

Lanelater fuscipes (Fabricius)

Elater fuscipes Fabricius, 1775 : 211.
Agrypnus fuscipes (Fabricius) Germar, 1840 : 253.
Lanelater fuscipes (Fabricius) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens from ‘India
orientali Koenig’. Koenig collected at Tranquebar in Southern India (Zimsen,
1964: 12). The two specimens in the UZM, Copenhagen, recorded by Zimsen
(1964 : 158), may be syntypes.

The generic attribution is based on material standing over this name in the
BMNH and MNHN, Paris.

Lanelater fusiformis (Candéze)

Agrypnus fusiformis Candéze, 1857 : 39.
Lanelater fusiformis (Candéze) Arnett, 1952 : 105.

Holotype. Curina: Chine meridionale, Mniszech collection from Buquet. The
material is in the IRSNB, Brussels according to Fleutiaux (1947: 290). Not
examined.

The confirmation of Arnett’s generic attribution is based on the description.

Lanelater gestroi (Candéze)

Agrypnus gestroi Candéze, 1880 : 188.
Lanelater gestyvoi (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Binaghi, 1941) : 88). NEw GUINEA: 3, Nuova Guinea,
Fly River 1876-77, legit L. M. D’Albertis (CMSN, Milan). Not examined.

The description is based on an unrecorded number of specimens. Binaghi
records only one specimen. He does not state whether it bears Candéze’s determina-
tion label.

The confirmation of Arnett’s generic diagnosis is based on the description.

Lanelater grandis (Hope)

Agrypnus grandis Hope, 1842 : 428.
Lanelatey grandis (Hope) Van Zwaluwenburg, 1959 : 350.

Lectotype (designated by Van Zwaluwenburg, 1959). AUSTRALIA: 9, grandis
Hope, Pt. Ess. [Hope] (UM, Oxford).

250 Cy MM. Fs vonesHAvEK

Lanelater gutturosus (Fairmaire)

Agrypnus gutturosus Fairmaire, 1884¢c : CX XIII.
Lanelatey gutturosus (Fairmaire) Arnett, 1952 : I05.

LECTOTYPE (present designation). Somartr REpuBLic: 9, Museum Paris,
Somali, Revoil 1883 [printed]; type; Agrypnus gutturosus n.sp. [Fairm.]; Agrypnus
gutturosus Fairm., type [Fleut.] (MNHN, Paris).

The description is based on an unrecorded number of specimens from Makidischu
[probably Mogadisco]. The specimen also bears a small label with an illegible
number which may correspond to an untraced list of localities. The printed
locality label was probably affixed as a matter of curatorial routine.

Lanelater hageni (Candéze)

Agrypnus hageni Candéze, 1887a : 189.
Lanelater hageni (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Van Zwaluwenburg, 1959 : 350). SUMATRA: Q, Dr B.
Hagen, Tandjong Morawa, Serdang (N. O. Sumatra); 1; A. Hageni Cdz. n.sp.
[Cand.] (RNH, Leiden).

Paralectotype. 9, locality as lectotype, without Candéze’s determination label.
Numbered ‘2’ by van Zwaluwenburg (RNH, Leiden).

The description is based on two specimens. The specimens numbered 3
($, H. M. Pantekok Bekri, Dali) and 4 (9, J. D. Pasteur, Insula Nias) by van
Zwaluwenburg (1959 : 350) are not part of the syntype series.

Lanelater infuscatus (Klug)

Agrypnus infuscatus Klug, 1855 : 67.
Agrypnus antennatus Candéze, 1893 : 5. [Synonymized by Schwarz, 1906 : 8.]
Lanelatey infuscatus (Klug) Arnett, 1952 : 105.

Agrypnus infuscatus Klug. LECTOTYPE (present designation). MOZAMBIQUE:
6, Tette, Peters Nr. 16047; infuscatus Klug, Cand. Tette, Peters [Gerstaecker]
(NMHU, Berlin). Length 19:5 mm.

The number 16047 refers to a museum catalogue recording the locality and
collector. The absence of Klug’s label is probably due to Gerstaecker (see p. 275).

Klug recorded the size range of this species as 84-9} [presumably German]
lines (=18-6-20-°8 mm). The NMHU, Berlin collection contains two specimens
with the same locality labels as the lectotype measuring 24mm in length. The
discrepancy in length is too great to allow these specimens to be accepted as part
of the original syntype series. They are conspecific with the lectotype.

Agrypnus antennatus Candéze. LECTOTYPE (present designation). SOUTH
AFRICA: g, n.sp. antennatus Cand., Natal, Per. [Cand., blue border]; A. antennatus;

Agrypnus antennatus Cand., det. E. Candéze [IRSNB curatorial label] (IRSNB,
Brussels).

RECLASSIFICZTION OF SUBFAMILY AGRYPNINAE 251

Lanelater javanus (Candéze)

Agrypnus javanus Candéze, 1857 : 44.
Lanelater javanus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). JAvA: 9, Dej. Java; Janson coll. 1963.130;
A. javanus [Cand.] Agrypnus javanus (Dej. Cat.) Cdze, Cand. type (e coll. Dejean)
[Janson] (BMNH).

Paralectotype. 9, Dej. Java; Janson coll. 1903.130; Agrypnus javanus (Dej.
Cat.) Cdze, Cand. (e coll. Dejean) [Janson] (BMNH).

Lanelater judaicus (Reiche & Saulcy)

Agrypnus judaicus Reiche & Saulcy, 1857 : 418; pl. 12, fig. 11.
Lanelater judaicus (Reiche & Saulcy) Arnett, 1952 : 105.

The description is based on material collected by Sauley at Jerusalem with
the comment ‘J’en ai vu un individu de Bayrouth et un autre de Saida’.
Neither of these specimens has been traced.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater labeculatus (Candéze)

Agrypnus labeculatus Candéze, 1892a : 795.
Lanelater labeculatus (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Binaghi, 1941b: 87). ENnGGaNo: Kifu-juc (CMSN,
Milan). Not examined.

Paralectotypes: 2 ex, Bua-Bua (CMSN, Milan). Not examined. ? 9, Engano,
Kifu-juc, v. Modligiani 1891; Agrypnus labeculatus Cand. n.sp. [?Cand.]; ex type,
Gestro det. [Fleut.]. ? 3, Engano, Bua-Bua, Modigliani 1891; Agrypnus labeculatus
Cand. [Cand.] (MNHN, Paris). The contents of the abdomen of each of the Paris
specimens has been eaten by Anthrenus.

The description is based on an unrecorded number of specimens (‘plusieurs
individus’) collected on Enggano at Bua-Bua and Kifu-juc in May and June 1891
by Modigliani and submitted to Candéze by Gestro. Binaghi does not record
the presence of Candéze’s determination label. The IRSNB, Brussels may possess
additional paralectotypes. The island of Enggano (or Engano) lies about 70 miles
off the west coast of southern Sumatra.

Lanelater lacertosus (Candéze)

Agrypnus lacertosus Candéze, 1857 : 38; pl. 1, fig. I.
Lanelater lacertosus (Candéze) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens with a size-
range of 30-40 mm from Silhet [=Sylhet, E. PaxIsTAn].

Syntype-material: not found in BMNH. ?IRSNB, Brussels.

252 C. M. F. von HAYEK

Confirmation of Arnett’s generic attribution is based on specimens standing as
lacertosus in the BMNH.

Lanelater laosensis Ohira
Lanelater laosensis Ohira, 1970b : 232; pl. VI, fig. C, pl. VII, fig. F.

Holotype. 3, Laos (TM, Budapest). Not examined.

Paratype. 9, Laos (TM, Budapest or Entomological laboratory, Aichi Uni-
versity, Japan). Not examined.

Lanelater laticollis (Hope) comb. n.
Agrvypnus laticollis Hope, 1843 : 366.
Agrypnus caliginosus Candéze, 1857 : 28. [Confirmation of Candéze’s (1857 : 28) provisional
synonymy. |
Lanelater caliginosus (Candéze) Arnett, 1952 : 105.

Agrypnus laticolis Hope. LECTOTYPE (present designation). W. AFRICA:
3, laticollis Hope Palmas [Hope]; Type, Hope, Ann. Nat. Hist. 11. 1843 p. 366.
Coll. Hope Oxon; Type coll. 1545, Agrypnus laticollis Hope. Hope Dept, Oxford
(UM, Oxford).

Agrypnus caliginosus Candéze. LECTOTYPE (present designation). SENEGAL:
3, Senegal Dej.; Janson coll. 1903.130; A. caliginosus [Cand.]; Agrypnus caliginosus
(Buq.) Cdze, Cand. type (A. senegalensis var. Dej. Cat. e coll. Dejean) [Janson]
(BMNH).

The absence of Dejean’s label is probably due to Janson (see p. 276). The
specimen is 38 mm long compared with the published length of 40 mm.

Schenkling (1925 : 7) records Jaticollis as a synonym of caliginosus. The Hope
name has priority.

Lanelater latior (MacLeay) sp. rev., comb. n.
Agrypnus latioy MacLeay, 1872 : 250.

The description is based on an unrecorded number of specimens collected by
Masters at Gayndah [AusTRALIA, Queensland]. Neboiss (1956: 3) states that
the type is in the AM, Sydney but since he does not record the data or otherwise
identify the specimen, this cannot be accepted as a valid lectotype designation.

Candéze (1874 : 13) synonymized this species with mastersi MacLeay. Neboiss
(1953 : 3) accepted the synonymy. The reason for treating Jatior as a valid species
is discussed under masters on p. 255.

The generic attribution is based on the description.

Lanelater longicollis (Candéze)

Agrypnus longicollis Candéze, 1889 : 63 (3).
Lanelater longicollis (Candéze) Arnett, 1952 : 105.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 253

The description is based on an unrecorded number of specimens from ABYSSINIA.
Type-material: ?IRSNB, Brussels.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater longicornis (Gahan) comb. n.
Agrypnus longicornis Gahan, 1900 : 26; pl. I, fig. 11.

Holotype. SoMALI REPUBLIC: g, Cent. and East Somaliland. Capt. June 5—
October 29.97 and pres. 1897 by C. V. A. Peel; Agrypnus longicornis Gahan. Type
[Gahan]; 1900.128 [presented by Prof. E. B. Poulton, Oxford] (BMNH).

The published locality is Central or East Somaliland.

This species is not listed in the main part of the Schenkling (1925) Catalogue
and is assumed to have been overlooked by Arnett (1952). See p. 241.

Lanelater lopezei (Fleutiaux)

Agrypnus lopezei Fleutiaux, 1934¢ : 477.
Lanelater lopezei (Fleutiaux) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens in the collections
of the author and the Hawaiian Sugar Planters’ Experimental Station, Hawaii.

Syntype examined. PuHILipPINEs: ¢g, Diklom, Bukidnon, Mindanao P. I.
2000 ft, 3.26.32 [altitude altered from 3000 ft and date added in manuscript].
F. C. Haddon collection; Agrypnus lopezei Fleut. type [Fleut.]. 1 3, same locality
but with the printed altitude 3000 ft (MNHN, Paris).

Lanelater luridus (Fabricius)

Elater luridus Fabricius, 1781 : 265.

Agrypnus punctatus Candéze, 1857 : 26. Syn. n.
Agrypnus sondaicus Candéze, 1857 : 33. Syn. n.
Agrypnus moestus Candéze, 1857 : 34. Syn. n.
Agrypnus giluus Candéze, 1865: 5. Syn. n.
Lanelater luridus (Fabricius) Arnett, 1952 : 105.

Elater luridus Fabricius. LECTOTYPE (present designation). dg, Elat. luridus
Fabr. sp. Ins. n. 3 [unidentified handwriting] (BMNH, Banks collection).

Paralectotype: 9, head and prothorax lost, no labels (BMNH, Banks collection).

The published locality is Coromandel [INp1A]. The paralectotype does not
agree with the females of the BMNH Juridus series and probably belongs to a
different species.

Agrypnus punctatus Candéze. LECTOTYPE (present designation). JAvA: 9,

Java; Janson coll. 1903.130; Agrypnus punctatus Cand., Cand., type (e coll.
Laferté); D. Hope (BMNH).

254 Cc. M. F. von HAYEK

Paralectotype: 3, Deyr.; Janson coll. 1903.130; Agrypnus punctatus Cdze, C and.
(e coll. Dyrolle) [Janson]; D. Hope (BMNH).

The absence of Candéze’s determination label is probably due to Janson (see
p. 276). Not asynonym of aequalis as stated by Fleutiaux (1889 : 138).

Agrypnus sondaicus Candéze. Holotype. JAvA: 9, Dej. Java; Janson coll.
1903.130; sondaicus [Cand.]. Agrypnus sondaicus (Dej. cat.) Cdze, Cand. type
(e coll. Dejean) [Janson] (BMNH).

Not a synonym of aequalis as stated by Fleutiaux (1923 : 409).

Agrypnus moestus Candéze. LECTOTYPE (present designation). InpIA: 4,
Dej. India; Janson coll. 1903.130; Agrypnus moestus Cdze. A socius Dej. coll.
(e coll. Dejean) [Janson] (BMNH). Length: 25 mm.

The description is based on an unrecorded number of specimens from ‘Des Indes
Orientales’ (see p. 271). Candéze lists Agrypnus socius Dej. Coll. [nomen nudum]
as a synonym. Despite the absence of Candéze’s determination label I believe
that this specimen formed part of the series on which he based his description.
The absence of Candéze’s determination label and the discrepancy between the
published locality and that on the label are probably due to Janson (see p. 276).

L. moestus (Candéze) is not conspecific with L. aequalis (Candéze) as recorded
by Fleutiaux (1889: 138) nor is the synonymy (Fleutiaux, 1926: 93) with
L. funestus (Candéze) correct.

Fleutiaux (1935 : 194) records luridus from Ethiopia, Bourillé, Kenya, Lokitang
and also Arabia, Somaliland, Abyssinia, Uganda and Tanganyika. The MNHN,
Paris contains the following specimens standing as Juridus in the collection: 2 J,
7 © Bourie [sic, see p. 260], 1 g, Lokitang, 1 3, Abyssinie, Harar, 1 g, Unguru
D.O.A. [=Tanganyika]. None of these specimens is conspecific with the type
of luridus and all belong to an unnamed African species. It seems probable that
luridus does not occur in Africa and that the distribution of this species is limited
to India and Ceylon.

Agrypnus giluus Candéze. The description is based on an unrecorded number
of specimens from S1AM and CAMBODIA received from de Castelnau.

Type-material: not found in the BMNH or the IRSNB, Brussels or Castelnau
collection, NMV, Melbourne. Interpretation of the species is based on the following
specimen. 4, Malacca; Janson coll. 1903 : 130; Agrypnus gilvus Cdz. Malacca
[Cand.]; moestus [Cand.] 1874 [Janson] Agrypnus gilvus 1864, A. moestus Cdze.
1874 ex coll. Candéze [Janson] (BMNH). This is the only specimen with Candéze’s
determination label known to me. Not a synonym of aequalis as stated by
Candéze (1874 : Io).

Lanelater maculicollis (Gerstaecker)

Agrypnus maculicollis Gerstaecker, 1871 : 53.
Lanelater maculicollis (Gerstaecker) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens with a size-
range of 22-25mm from ZANZIBAR collected by van der Decken. Candéze

|

RECLASSIFICATION OF SUBFAMILY, AGRYPNINAE 255

(1874 : 7) records that the type is in the ZMHU, Berlin (not confirmed). Interpre-
tation of the species is based on the three specimens (2 2, I g) from MozAMBIQUE
recorded by Candéze (1874 : 7) in the BMNH.

Lanelater mastersi (MacLeay)

Agrypnus mastersi Macleay, 1872 : 249.
Lanelater mastersi (Macleay) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens collected by
Masters at Gayndah [AusTRALIA, Queensland]. Neboiss (1956: 3) records that
the type is in the AM, Sydney, but since he does not record data or otherwise
identify the specimen this statement cannot be accepted as a valid lectotype
designation.

Candéze (1874 : 13) believed mastersi to be a very variable species and regarded
A. lator MacLeay as a synonym. Neboiss (1953: 3) accepted the synonymy.
However, since the specimens standing as mastersi in the BMNH belong to a
number of different species, suggesting that mastersi is not as variable as Candéze
supposed, I consider it prudent to treat Jatvor MacLeay as a distinct species until
the lectotypes of the two species have been designated and compared.

Lanelater modestus (Schwarz)

Agrypnus modestus Schwarz, 1902f : 305.
Lanelatey modestus (Schwarz) Arnett, 1952 : 105.

Holotype. LrssErR SunpA IsLANDs: Sex unknown, Sumbawa (Moser) ?DEI,
Eberswalde.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater moseri (Schwarz)

Agrypnus moseri Schwarz, 1903a : 357.
Lanelater moseri (Schwarz) Arnett, 1952 : 105.

Syntypes: TANZANIA: 2 ex., Deutsch Ostafrica (Moser). ? DEI, Eberswalde.
Confirmation of Arnett’s generic attribution is based on the description.

Lanelater mucronatus (Candéze)

Agrypnus mucronatus Candéze, 1857 : 42.
Agrypnus mucronatus var. incisus Fleutiaux, 1927 : 59.
Lanelater mucronatus (Candéze) Arnett, 1952 : 105.

Holotype. Borneo: g, mucronatus Cdz. Borneo [Candéze, yellow border];
Agrypnus mucronatus Cand. det. E. Candéze [IRSNB curatorial label]; Collection
E. Candéze (IRSNB, Brussels).

256 C. M. F. von HAYEK

A. mucronatus var incisus Fleutiaux. Material examined: 1 9, Luang Prabang,
Environs. Vitalis de Salvaza; Bou Nam Mo, Luang Prab. 3.3.18 mucronatus Cand.
v. incisus Fleut. [Fleut.] (MNHN, Paris).

Lanelater nitidus (Fleutiaux)

Agrypnus nitidus Fleutiaux, 1918c : 167.
Lanelatey mtidus (Fleutiaux) Arnett, 1952 : 105.

LECTOTYPE (present designation). KENyA: 9, Kedong Valley, B.E.A.,
G.B. Mars 1913; Agrypnus nitidus Fleut., type [Fleut.] (MNHN, Paris). The
Kedong Valley is about 35 miles east of Nairobi.

Lanelater notodonta (Latreille)

Elatey notodonta Latreille, 1827 : 275; pl. 58, fig. 6.

Agrypnus notodonta (Latreille) Germar, 1840 : 253.

Amaurus senegalensis Castelnau, 1840 : 237. [Synonymized by Candéze, 1857 : 27.]
Agrypnus himerensis Ragusa, 1881 : 8; pl. I, figs 7 & 8.

Agrypnus notodonta var. himerensis Ragusa, Ragusa, 1911 : 196.

Agrypnus notodonta ssp. imerensis Binaghi, 1941a : 72. [Unjustified emendation. ]
Lanelatey notodonta (Latreille) Arnett, 1952 : 105.

Elater notodonta Latreille. The description is based on an unrecorded number
of specimens from Sennar [SUDAN] collected by Caillaud in the course of his
expedition to Meroé. There are no specimens from the Latreille collection in the
BMNH. The location of the Caillaud collection is unknown.

Interpretation of the species is based on the following specimen: Ecypt: 9,
Egypt; Janson coll. 1903.130; Agrypnus notodonta Latr, Egypte [Cand.]; Agrypnus
notodonta Latr. Cdze. ex coll. Candéze [Janson] (BMNH). Examination of the
available material standing as notodonta in the BMNH, IRSNB, Brussels and
MNHN, Paris has shown that there is no disagreement concerning the identity
of this species.

Amaurus senegalensis Castelnau. The description is based on an unrecorded
number of specimens from SENEGAL. The material cannot be found in the IRSNB,
Brussels. The interpretation of the species is based on the following specimens:
2 2 on one cork mount, notodonta (Senegal) (NMV, Melbourne, Castelnau collection).
These specimens may be the ones on which Castelnau based his description and
which have been relabelled at a later date. They agree well with the description
and measure 28-5 mm and 28 mm in length and 9 mm and 8 mm in width compared
with the published measurements of 14 lines [= 31-5 mm (French lines) or 29-6 mm
(English lines)] and 4 lines [= 9 mm or 8-4 mm].

Agrypnus himerensis Ragusa. The description is based on an unrecorded number
of specimens collected in Siciry, Termini Imerensi in August and September.
Ragusa first saw the species standing as Agryphus conspersus Dej. in the Baldassare
Romano collection [not located by the present writer] in 1870 and again in the

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 257

collection of Prof. Ciafalo from whom he received six specimens. The Ragusa
collection was broken up (Horn & Kahle, 1936: 218) and it is therefore possible
that the following specimens are part of the original syntype-series.

Material examined; 1 3, himerensis Rag. Sicil.; Museum Paris, Collection Leon
Fairmaire 1906. 1 9, Type, Sicile, Ragusa; Museum Paris, Coll. L. Bedel 1922;
himerensis Rag. 1 9, Sicile, Termini, E. Ragusa; Termini; Agrypnus himerensis;
himerensis Rag. [Fleut] (MNHN, Paris). The handwriting on the determination
labels is unknown and appears to be different in each case.

Schenkling (1925 : 4) records the following species as a synonym of notodonta
Latreille.

Elater fuscipes Olivier, 1790; no. 31, p. 20; pl. 3, fig. 21. The description is
based on an unrecorded number of specimens from ‘Cap de Bonne-Esperance’
S. Africa. Fleutiaux (1911 : 475) synonymized fuscipes with notodonta, commenting
that Olivier’s type is in his collection. However the male in the Fleutiaux collec-
tion in the MNHN, Paris labelled ‘fuscipes Ol. non Fab., type d’Olivier’ by Fleutiaux
bears the locality SENEGAL. It is notodonta Latreille. L. notodonta has not been
recorded from S. Africa. Agvypnus antennatus Candéze (= L. infuscatus (Klug)),
and L. peringueyr Candéze are the only Lanelater species recorded from S. Africa.

Lanelater ocellatus (Candéze)

Agrypnus ocellatus Candéze, 1857 : 39.
Lanelater ocellatus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). ¢, Ind. or.; Janson coll. ex Candéze,
1903.130; Agrypnus ocellatus Cdz., Ind. or. [Cand.]; Agrypnus ocellatus Cand.,
type. Ex coll. Cand. [Gahan] (BMNH).

The published locality is ‘Des Indes Orientales’ (see p. 271).

Additional material examined. CEYLON: Colombo, on coast level, 7-27.iv.32
(G. Lewis) 1 g,19 (BMNH).

Lanelater olcesii (Fairmaire)

Agrypnus olcesit Fairmaire, 1884a : 446.
Lanelater olcesi1 (Fairmaire) Arnett, 1952 : 105.

LECTOTYPE. Morocco: 92, Mogador; Agrypnus olcesii Frm. Mar. [?, illegible]
1884 [Fairm.] Type; Museum Paris, Coll. L. Fairmaire (MNHN, Paris).

Lanelater opacus (Candéze)

Agrypnus opacus Candéze, 1878 : 8.
Lanelater opacus (Candéze) Arnett, 1952 : 105.

| LECTOTYPE (present designation). Wrst Matraysia: g, Malacca; Janson
| coll. 1903.130; A. opacus Cdz., type, Malacca [Cand.]; Agrypnus opacus Cdze,
| type ex coll. Castelnau [Janson] (BMNH).

R

258 cM. BF. von HAYEK

Paralectotype: 9, Malacca; Janson coll. 1903.130; Agrypnus opacus Cdz, Cand.,
ex coll. Castelnau [Janson] (BMNH).

Lanelater pacificus (Candéze)

Agrypnus pacificus Candéze, 1882 : I.
Lanelater pacificus (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Van Zwaluwenburg, 1959: 350, see p. 9). NEW
GUINEA: 9 Woodlark I. (IRSNB, Brussels). Not examined. Van Zwaluwenburg’s
comment ‘designated by Candéze’ presumably indicates that the specimen bears
Candéze’s determination label with word ‘type’.

Candéze did not record this species from New Britain. Therefore the two addi-
tional specimens in the IRSNB Brussels to which van Zwaluwenburg refers, cannot
be part of Candéze’s original series.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater parallelicollis (Candéze)

Agrypnus parallelicollis Candéze, 1889 : 63 (3).
Lanelater parallelicollis (Candéze) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens, with a size-
range of 20-30 mm, from Kordofan [SuDAN].
Type-material: ? IRSNB, Brussels.

The generic attribution is based on a male from N. NicEeriA, Anzare, determined
by Fleutiaux (BMNH).

Lanelater peringueyi (Candéze)
Agrypnus peringueyi Candéze, 1889 : 68 (2).
Lanelater peringueyi (Candéze) Arnett, 1952 : 105.
The description is based on an unrecorded number of specimens from Cape
Town [S. AFRICA] received from Peringuey.
Type-material: ? IRSNB, Brussels.
Confirmation of Arnett’s generic attribution is based on the description.

Lanelater permucronatus (Schwarz)

Agrypnus permucronatus Schwarz, 1902) : 194.
- Lanelater permucronatus (Schwarz) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens, with a size-
range of 28-33 mm, from Borneo, Kina Balu.

Syntype-series: ? DEI, Eberswalde.
Confirmation of Arnett’s generic attribution is based on the description.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 259

Lanelater persicus (Candéze)

Agrypnus persicus Candéze, 1874 : 6.
Lanelater persicus (Candéze) Arnett, 1952 : 105.

Holotype. Persia: g, 39865 [Fry catalogue number = Persia, Millingen
(Higgins) ] Persia; Fry coll.1905.100; A. persicus Cdz., type [Cand.]; Agrypnus persicus
Cand., type ex coll. Fry [Gahan]; Agrypnus persicus Cand., type [Fry] (BMNH).

Lanelater pescadoriensis (Miwa)

Agrypnus pescadoriensis Miwa, 1934 : 178; pl. 9, fig. 9.
Lanelater pescadoriensis (Miwa) Arnett, 1952 : 105

The description is based on an unrecorded number of specimens from Hokoto
(Pescadores), V. 1932 (T. Chin).

Type-material: ?Taiwan Agricultural Research Institute, Taipei.
Confirmation of Arnett’s generic attribution is based on the description.

Lanelater politus (Candéze)

Agrypnus politus Candéze, 1857 : 43.
Lanelatey politus (Candéze) Arnett, 1952 : 105.

Holotype. CHINA: 9, Gory, China; Janson coll. 1903.130; politus Gory China
[? Gory]; A. politus [Cand.]; Agrypnus politus (Gory) Cdze., Cand., type (e coll.
de Laferté) [Janson] (BMNH).

Lanelater ponderatus (Candéze)

Agrypnus ponderatus Candéze, 1897 : 5.
Lanelater ponderatus (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Van Zwaluwenburg, 1959: 350). PHILIPPINES:
? 9, n.sp. ponderatus Cand. Mindanao St.faudinger] [presumably Cand.] (IRSNB,
Brussels).

Paralectotype: second specimen, data not recorded, listed by Van Zwaluwenburg
(1959 : 350) (IRSNB, Brussels).

Lanelater proximus (Fleutiaux)

Agrypnus proximus Fleutiaux, 1935) : 194.
Lanelater proximus (Fleutiaux) Arnett, 1952 : 105.

LECTOTYPE (present designation). Etuiopia: 9, Ethiopia Merid. Bourie
[sic] Bord de la Riv. Omo, 600m; Agrypnus proximus Fleut., type, gj [Fleut]
(MNHN, Paris).

Paralectotype. KENyA: 9, Bords du lac Rodolphe, Monts Lubur, 570 m;
proximus Fleut., type 9 [Fleut] (MNHN, Paris).

260 C. M. F. von HAYEK

The published locality is Bourillé, an area on the west bank of the river Omo,
about 40 miles north of the northern end of Lake Rudolph. Bourie is a misprint
which occurs on a large number of labels. Fleutiaux was mistaken in the sex of
the specimen from Bourillé.

Lanelater puber (Candéze)
Agrypnus puber Candéze, 1857 : 30.
Lanelater pubey (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). SENEGAL: 9, Senegal, Dej.; Janson coll.
1903.130; puber [Cand.]; Agrypnus puber (Dej. Cat.) Cdze, Cand, type (e coll.
Dejean) [Janson] (BMNH).

Paralectotypes. 1 9, 2 g, Dej. Senegal: Janson coll. 1903.130; Agrypnus puber
(Dej. Cat.) Cdze, Cand. (e coll. Dejean) [Janson] (BMNH).

A. tropicus Hope (1843 : 365), which Candéze tentatively recorded as a synonym,
is a Propsephus species (see p. II).

Lanelater pubescens (Candéze)
Agrypnus pubescens Candéze, 1857 : 31.
Agrvypnus bocandei Candéze, 1857 : 32. [Reduced to a variety of pubescens Candéze, Candéze,

DOT ANAS
Lanelatey pubescens (Candéze) Arnett, 1952 : 105.

Agrypnus pubescens Candéze. LECTOTYPE (present designation). SENEGAL:
9, Senegal; Janson coll. 1903.130; A. pubescens Cdz. 1 (p. 31) [Cand.]; Agrypnus
pubescens Cand. v. [Janson] (BMNH).

Candéze mentions both sexes in his description, but no male syntypes have
been located. The lectotype agrees with the description but measures 20 mm in
length compared with the published measurements of 25mm. As there is a similar
discrepancy in the case of bocandi it has been assumed that Candéze made a 5 mm
error when measuring these specimens.

Agrypnus bocander Candéze. LECTOTYPE (present designation). SENEGAL:
6, Senegal, Dej.; bocandei [Cand.]; Agrypnus bocandei (Dej. coll.) Cdze, Cand.,
type (e coll. Dejean) [Janson] (BMNH).

The specimen measures 17-2 mm in length compared with the published length
of 12mm. This discrepancy is discussed above. Fleutiaux (1935): 195) con-
sidered that bocandei should be regarded as a good species. Comparison of the
lectotypes has shown that this belief is not justified.

Lanelater pumilus (Candéze)

Agrypnus pumilus Candéze, 1889 : 69 (3).
Lanelater pumilus (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Binaghi, 1941): 89). Etuiopia: Assab, IV, 1888,
legit Ragazzi. (CMSN, Genoa). Not examined.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 261

The description is based on an unrecorded number of specimens in the MCSN,
Genoa. Binaghi records only one specimen. He does not state whether it bears
Candéze’s determination label.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater renardi (Candéze)
Agrypnus venardi Candéze, 1890 : CXLVIII.

Lanelater venardi (Candéze) Arnett, 1952 : 105.
The description is based on 1 ex. from Tetara and 2 ex. from Konbir [INnp1A]
collected by Cardon (see p. 273).

Syntypes: ? IRSNB, Brussels.
Confirmation of Arnett’s generic attribution is based on the description.

Lanelater resectus (Candéze)

Agrypnus resectus Candéze, 1857 : 45.
Lanelater vesectus (Candéze) Arnett, 1952 : 105.

The description is based on a single specimen from Nouvelle Hollande
[AUSTRALIA] bearing the name Agrypnus australasiae Dup. in the Mniszech
collection.

Holotype: IRSNB, Brussels according to Neboiss (1961 : 5). Not confirmed.

Confirmation of Arnett’s generic attribution is based on specimens standing
as vesectus in the BMNH.

Lanelater robustus (Fleutiaux)

Agrypnus vobustus Fleutiaux, 1902a@ : 163.
Lanelater vobustus (Fleutiaux) Arnett, 1952 : 105.

LECTOTYPE (present designation). JAvaA: gj, Java, Agrypnus robustus Fleut.,
type [Fleut.] (MNHN, Paris).

Paralectotypes. I ex., Java. 6 3, 5 9, Java occident. Pengalengen, 4000,
1893. H. Fruhstorfer. One specimen bears Fleutiaux’s determination label.
I dg, 1 9, Palaboean, Java, S.O. 1 4, 3 9, Soekaboemi, Java, Rouyer (MNHN,
Paris). These 18 specimens stood beside Fleutiaux’s type-specimen in the Fleutiaux
collection. Since they fall within the published size-range of 38-45 mm, it seems
safe to assume that Fleutiaux had them before him at the time of the description.

| Lanelater rubiginosus (Candéze)
| Agrypnus rubiginosus Candéze, 1865 : 5.
Lanelater rubiginosus (Candéze) Arnett, 1952 : 105.

| Lectotype (designated by Van Zwaluwenburg, 1959 : 351). SUMATRA: 9, Miiller,

262 Cris ES wont EA EK

Sumatra, presumably with Candéze’s determination label (RNH, Leiden). Not
examined.

Since Van Zwaluwenburg does not record the data on the second specimen it
is impossible to determine whether it formed part of the syntype-series.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater rufipes (Candéze)

Agrypnus rufipes Candéze, 1874 : 7.
Lanelater rufipes (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). Inp1A: 9, Pondicherty; Janson coll. 1903.
130; A. rufipes Cdz. n.sp. [Cand.]; Agrypnus rufipes Cdz., type [Janson] (BMNH).

Lanelater rufus (Candéze)

Agrypnus rufus Candéze, 1857 : 32.
Lanelater rufus (Candéze) Arnett, 1952 : 105.

LECTOTYPE (present designation). 9, type; Cayenne, typ.; Collection Chevrolat;
rufus Cand., type mon. [Fleut.]. The specimen stands over a green Chevrolat
label: Agrypnus rufus (Chev.) Candéze type. Mon. 1, p. 32. 13. Cayenne, Coll.
Chevrolat (MNHN, Paris).

The published locality is ‘Guyane’ but Candéze (1874: 6) states that this is
erroneous and that rufus is an African species. It has been recorded as an African
species in subsequent catalogues (Candéze, 1891: 10, Schenkling, 1925: 7) but
there are no published records of its capture in Africa.

Lanelater sallei (LeConte)

Agrypnus sallet LeConte, 1853 : 491.
Lanelater sallei (LeConte) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens with a size-
range of 0-85-1-10 [inches = 21-7—28-:2 mm.] collected near New Orleans [U.S.A.,
Louisiana] by Sallé and given to LeConte by Guex and specimens subsequently
collected by Schott at Eagle Pass [Texas] on the Lower Rio Grande.

LECTOTYPE (present designation). U.S.A.: 3, brick red paper disk [Southern
States, see p. 277]; Type, 2373 [MCZ curatorial label]; Agrypnus sallei Lec. [LeC.]
(MCZ, Harvard). Length: 24-8 mm.

Paralectotypes: I 3, dark red paper disk [Texas, see p. 277]; sallei 3 [MCZ cura-
torial label]. Length: 21:8mm. 1 4, dark red paper disk [Texas]; sallei 4 [MCZ
curatorial label]. Length: 27-7 mm. (MCZ, Harvard). It is by no means certain
that these two specimens were in fact collected by Schott at Eagle Pass but since
they are the only specimens from Texas in the LeConte collection there is no
alternative but to accept them as part of LeConte’s syntype series. The two

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 263

paralectotypes are not conspecific with one another. The one labeled ‘sallei 3’
belongs to a species unknown to me. It is conspecific with two males from Brewster
county, Texas in the CAS, San Francisco collection. The specimen marked ‘sallei
4° differs from the lectotype in that the antennae do not attain the posterior angles
of the prothorax, falling short of it by the length of more than one segment. In
addition the lateral lobes of the aedeagus are stouter than those of the lectotype.
The specimen is conspecific with the males of series from Falfurrias, Texas, Green
[?Greer] Co., Oklahoma and Medora, Kansas. Further studies are required before
it can be decided whether these specimens belong to a distinct species or whether
they are a form of sallev.

Additional material examined. U.S.A., Louisiana: New Orleans, 1 9 (BMNH).
Florida: Dunedin, 2 ¢; Enterprise, 3 9; Hillsbro, 1 g, 1 9; Jacksonville, 1 9; Lake
Placid, r g; Naples, 1 g; St. Augustine, xr g; Stuart, 1 g (CAS, San Francisco.)
I have not seen any specimens of sallei from localities west of New Orleans.

The specimen labelled ‘sallei 2’ (3, grey-green paper disk [?faded dark green =
New Mexico, see p. 277] which is conspecific with specimens from Tucson believed
to be arizonae Candéze), and ‘Sallei 5’ (g. Fla., correctly identified) standing
with the syntypes in the LeConte collection (MCZ, Harvard) are obviously not
part of the syntype series.

Lanelater schotti (LeConte)

Agrypnus schotti LeConte, 1853 : 492.
Lanelater schotti (LeConte) Arnett, 1952 : 105.

Holotype. U.S.A.: 9, dark red paper disk [Texas, see p. 277]; Type 2374 [MCZ
curatorial label} (MCZ, Harvard). The published locality is Lower Rio Grande,
the river which forms the boundary between Texas and Mexico.

The LeConte collection (MCN, Harvard) also contains the following specimens:
I Q, illegible locality, 25; schotti 2 [MCZ curatorial label]. 1 9, Ariz.; schotti 3
[MCZ curatorial label]. These specimens, which are presumably those recorded
by Arnett (1952: 106), differ from his description in that the antennal grooves
do not have abrupt ends but become shallow at the base. In spite of the fact
that these two specimens have a more robust appearance, I believe that they are
conspecific with the holotype.

Lanelater scortecci (Binaghi) comb. n.
Agrypnus scortecci Binaghi, 1941a : 73; fig. 8.
Lanelater scortecci (Binaghi) Arnett, 1952 : 105.

Holotype and paratype. Lispya: Fezzan; Gat estate, 1934, light, G. Garganese
(MCSN, Milan) and another paratype, same data in Binaghi collection. Not
examined.

Confirmation of Arnett’s generic attribution is based on the description.

264 C>M. FP. von HAYEK

Lanelater semicribrosus (Fairmaire) sp. rev., comb. n.

Corymbites semicribrosus Fairmaire, 1887 : 153.
Agrypnus semicribrosus (Fairmaire) Candéze, 1896 : 6.
Agrypnus candezei Fleutiaux, 1919: 8. Syn.n.

Corymbites semicribrosus Fairmaire. LECTOTYPE (present designation).
SOMALI: §, Museum Paris, Somali, Revoil, 1885 [printed]; 490; [mss.]; Corymbites
semicribrosus Frm. [Fairm.]; Fairm. det. [Fleut.] (MNHN, Paris).

The published locality is Zanzibar. Fairmaire based his description on material
collected by Revoil during his expeditions to ‘les Somalis et dans l’interieur du
Zanguebar [see p. 44]. It seems probable that the printed label was attached
to all the material collected on these expeditions, and that the number 420 refers
to a more detailed list of localities whose present whereabouts is unknown.

Corymbites semicribrosus Fairmaire is not conspecific with Agrypnus infuscatus
Klug (see p. 250) as stated by Schwarz (1906 : 8).

Agrypnus candezei Fleutiaux. Holotype. TANzANIA: 9, Mandera, Zanguebar;
Agrypnus candezei Fleut., type [Fleut.]; 2 du preced. semicribrosus, Cand. det.
[Fleut.] (MNHN, Paris). The specimen has a broader, stouter appearance and
shorter, less strongly serrate antennae than most other female Lanelater species
known to me.

This synonymy is based on the belief that semicribrosus, of which the lectotype
resembles the males of a series of an undescribed species from Chiromo (Malawi)
in the BMNH, also resembles the undescribed species in that the females have a
broader, stouter appearance, and shorter, less strongly serrate antennae. This
belief is in some measure supported by the fact that a male with the same locality
label as the holotype of candezei agrees well with the lectotype of semuzcribrosus.
Fleutiaux did not publish the synonymy recorded on the holotype of candezev.

Lanelater semistriatus (Schwarz)

Agvypnus semistriatus Schwarz, 1899 : 74.
Lanelater semistriatus (Schwarz) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens with a size-
range of 22-23 mm from Deutsch Ost Africa [TANZANIA] collected by Bennigsen.

Syntype series: ? DEI, Eberswalde.

Confirmation of Arnett’s generic attribution is based on the description.

Lanelater sobrinus (Candéze)

Agrvypnus sobrinus Candéze, 1887a : 190.
Lanelater sobrinus (Candéze) Arnett, 1952 : 105.

Lectotype (designated by Van Zwaluwenburg, 1959 : 351). SUMATRA: 9, H. v.
Hasselt, Boenga Maas. Palembang, Sumatra; A. sobrinus Cdz. n.sp. [Cand.] (RNH,
Leiden).

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 265

Paralectotype: 2, Dr B. Hagen, Tandjong Morawa, Serdang (N.O. Sumatra);
A. sobrinus Cdz. A. sobrinus Cdz. n.sp. [Cand.]; 2 [Van Z.] (RNH, Leiden).

The second paralectotype, from the same locality as the lectotype has not been
located.

Lanelater soricinus (Candéze)

Agrypnus soricinus Candéze, 1882 : I.
Lanelater sovicinus (Candéze) Arnett, 1952 : 102.

The description is based on an unrecorded number of specimens from NEw GUINEA
near Gelwink Bay in the Lansberge and Candéze collections.

Syntype examined: 1 g, Cote Sept. 136.30.137 [°E. = Geelvink Bay]; soricinus
Cdz. n.sp. [Cand.]; Ex Museo Van Lansberge; Museum Paris. Collection Oberthur
(MNHN, Paris).

The IRSNB, Brussels may possess additional syntypes.

Lanelater substriatus (Candéze)

Agrypnus substriatus Candéze, 1857 : 25.

Agrypnus extractus Fleutiaux, 1902a : 162. [Synonymized by Schwarz, 1906 : 7.|
Agrypnus extractus var striatus Fleutiaux, 1902a : 162.

Lanelatey substriatus (Candéze) Arnett, 1952 : 105.

Agrypnus substriatus Candéze, LECTOTYPE (present designation). SENEGAL:
g, Senegal; Janson coll. 1903.130; substriatus [Cand.]; Agrypnus substriatus Cand.,
Cand., type (e coll. de Laferté) [Janson] (BMNH).

Paralectotype: 1 2, Senegal, Janson coll. 1903.130; Agrypnus substriatus Cand.
Cand. v [id] (e coll. de Laferté) [Janson] (BMNH).

The material from the Deyrolle collection has not been located.

Agrypnus extractus Fleutiaux. LECTOTYPE (present designation). CAMEROUN:
g, Cameroun; extractus Fleut., type [Fleut.] (MNHN, Paris).

Agrypnus extractus var. striatus Fleutiaux. LECTOTYPE (present designation).
SAN THOME: 9, San Thomé; extractus v. striatus Fleut., [Fleut.] (MNHN, Paris).

Lanelater tippooi (Candéze)

Agrypnus tippooi Candéze, 1889 : 69 (3).
Lanelater tippooi (Candéze) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens from INDIA:
Deccan méridional, Koimbatour.
Type-material: ?IRSNB, Brussels.

Confirmation of Arnett’s generic attribution is based on the description.

266 C.. Mo Be Von EEA ENS

Lanelater tomentosus (Fabricius)

Elater tomentosus Fabricius, 1798 : 138.
Agrvypnus tomentosus (Fabricius) Eschscholtz, 1829 : 32.
Lanelater tomentosus (Fabricius) Arnett, 1952 : 105.

The description is based on an unrecorded number of specimens from ‘India
orientali’. According to Zimsen (1964 : 158, no. 2612) this material has not been
located.

Confirmation of Arnett’s generic attribution is based on specimens standing as
tomentosus in the BMNH.

OCTOCRYPTUS Candéze

Octocryptus Candéze, 1892c : 486. Type-species: Octocryptus cardoni Candéze, by original
designation.

At first sight the five species included in the genus bear a very close resemblance
to the true Agrypninae. Examination of the underside immediately reveals the
distinctive characteristics of the group, the deep grooves near the lateral margins
of the propleurae for the accommodation of the antennae, and the fact that in
repose the tarsi are lodged in grooves occupying the position of the posterior portion
of the prosternopleural suture.

The relationship of this genus to other genera within the family is extremely
uncertain. Candéze erected the tribe Octocryptites for the genus. Schenkling
(1925 : 39) raised it to subfamily status while Fleutiaux (1941c) treats it as a sub-
division (Octocryptitae) of the Agrypninae.

Fleutiaux (1944 : 145) gives a generic diagnosis and key to species. In addition
to the diagnostic characters mentioned by Fleutiaux, the genus possesses the follow-
ing features: the mesepimera do not attain the margin of the middle costal cavity,
tibial spurs are absent and there are no setae at the base of each claw.

Fleutiaux (1944: 145) states that he examined the types of the two species
described by Candéze. However as Candéze did not record the number of specimens
on which he based his descriptions and as Fleutiaux provides no means of identify-
ing the specimens he examined, his statement cannot be accepted as valid lectotype
designations for cardoni Candéze and radula Candéze.

DISTRIBUTION. Oman, India, North Vietnam, Sumatra.

BIOLOGY AND HABITS. Nothing is known of the life history of the species.
Specimens have been found buried at some depth in leaf detritus near a river
(Candéze, 1892c : 487) and near lakes, streams and in damp places (Fleutiaux,
1944 : 148).

Octocryptus babaulti Fleutiaux

Octocryptus babaulti Fleutiaux, 1944 : 148.

LECTOTYPE (present designation). N.Inp1A: Sex unknown. Bajaura, Kangra
district (Indes Angl.); G. Babault, Juin 1914; boursouflures laterales du pronotum

=“

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 267

[illegible word] de sillons tar. sur des propleurs; Octocryptus babaulti Fleut., type
[Fleut.] (MNHN, Paris).

Octocryptus cardoni Candéze
Octocryptus cardoni Candéze, 1892c : 486.

The description is based on five specimens captured by Cardon in July on the
banks of the Sunk River in the Manguiers valley near Chichuani in Barwai (see
note on Cardon’s localities on p. 273).

Type-material: IRSNB, Brussels according to Fleutiaux, 1944 : 146. Inpma:
I ex., Barwai; Collection Candéze; Octocryptus cardoni Cand. det. E. Cand. [IRSNB
curatorial label]; cf. Ann. Soc. Ent. Belge VI, 1892 p. 487; ex typus; cardoni Cand.
Mus. Bruxelles, echange 1943 [Fleut.] (MNHN, Paris).

Octocryptus coomani Fleutiaux

[Octocryptus cardoni Candéze; Fleutiaux, 1927 : roo, fig. Misidentification. }
Octocryptus coomani Fleutiaux, 1944 : 148.

LECTOTYPE (present designation). NortH VIETNAM: 3, Tonkin, Lac Tho,
A. de Cooman; O. coomani Fleut., type, cardoni Fleut. non Cand. [Fleut.] (MNHN,
Paris).

Paralectotypes: 2 ex., same locality as the lectotype one with a note: a terre
dans le voisinage des etangs, ruisseaux, terrains humides en compagnie de Compso-
lacon. I 9, same locality as the lectotype: exemplaire figuré par Jeannel [probably
in Fleutiaux, 1942, Revue. fr. Ent. 9: 79 as cardoni|; Octocryptus coomani Fleut.
[Fleut.]. 1 9, same locality as the lectotype; type figure Faun. col. Franc. 1927
p. 100; cardoni Fleut. pars 1927, sp.? [Fleut.] (MNHN, Paris).

Octocryptus maindroni Fleutiaux
Octocryptus maindroni Fleutiaux, 1944 : 148.

LECTOTYPE (present designation). OMAN: 9, Mascate [= Muscat or Musquat|]
(Sept. Oct.) Maindron 133-96; Octocryptus maindroni Fleut., type [Fleut.] (MNHN,
Paris).

Octocryptus radula Candéze

Octocryptus radula Candéze, 18936 : 14.

The description is based on an unrecorded number of specimens from SUMATRA,
Padang.

Type-material: IRSNB, Brussels according to Fleutiaux, 1944: 147. Not
examined.

268 Ca Me BE. Von tEAM KG

CHECKLIST OF THE GENERA INCLUDED IN THE AGRYPNINAE
ADELOCERA Latreille EIDOLUS Candéze.

[LACON sensu Germar nec Castelnau.
Pars. ] OPATELUS Candéze.

AGRAEUS Candéze. Syn. n.

PERICUS Candéze. Syn. n.

BRACHYLACON Motschulsky. eet try aes

Syn. n. CRY PTOTARSUS Philippi nec
TRACHYLACON Motschulsky. Kirsch.
CAVICOXUM Pic. HEXAULACUS Candéze.

PROLACON Fleutiaux. Syn. n.

OS a ee DILOBITARSUS Lattreille.

SCAPHODERUS Candéze ANACANTHA Solier. Syn. n.

BRUYANTIUS Fleutiaux.

Syn. n. ELASMOSOMUS Schwarz.
LACON Castelnau HEMICLEUS Candéze.
[ADELOCERA sensu auct. nec
Latreille. ] AGRYPNUS Eschscholtz.

LEPIDOTUS Stephens nec Asso.
OCNEUS Candéze. Syn. n.
SCELISUS Candéze. Syn. n.
ALAOTY PUS Schwarz.
SULCILACON Fleutiaux. Syn. n.
DIPHYAULON Arnett. Syn. n.
AULACON Arnett. Syn. n.
ZALEPIA Arnett. Syn. n.
KOBULACON Chujo & Ohira.

Syn. n.

[LACON sensu Germar nec
Castelnau. Pars. ]
MECYNOCANTAUS Hope.
Syn. n.
TYLOTARSUS Germar. Syn. n.
TILOTARSUS Candéze.
MYRMODES Candeze. Syn. n.
ARCHONTAS Goezis.
PSEUDOLACON Blackburn.

, Syn. n.
LEPIDELATER Smith. Syn. n.
ARNETTIA Golbach. Syn. n. pace, po a

MONOCYRTON Golbach. Syn. n.
CORNILACON Golbach. Syn. n.
LATILACON Golbach. Syn. n.

LOBOTARSUS Schwarz. Syn. n.
LOBITARSUS Fleutiaux.
ENOPLODERES Schwarz nec

Falderman.
CANDANIUS nom. n. CENTROSTETHUS Schwarz.
ANIUS Candéze nec Pascoe. Syn. n.

COM PSOLACON Reitter.

PARALACON Reitter. Syn. n.

DANOSOMA Thomo). NEOLACON Miwa. Syn. n.
DELOX Quelle. COLAULON Arnett. Syn. n.

t=

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 269

CRY PTOLACON Nakane & RISMETAUS Fileutiaux.
Kishi. Syn. n.
SABIKIKORIUS Nakane & Kishi.
Syn. n. TRIERS Candéze.
SAGOJYO Kishi. Syn. n.
SAGOJO Ohira.

P@eONTOIDES Cobos, “Synia/ a (LANELATER Arnett.

PYRGANUS Golbach. Syn. n. [AGRYPNUS sensu auct. nec
Eschscholtz. |
MERISTHUS Candéze. AMAURUS Castelnau nec
Burmeister.

Subgenus MERISTHUS Candéze.
RHACIASPIS Arnett.
Subgenus SULCIMERUS Fleutiaux. OCTOCRYPTUS Candéze.

ACKNOWLEDGEMENTS

It gives me great pleasure to thank my colleague Mr R. D. Pope for his
encouragement, help, advice and apparently inexhaustible patience during the
preparation of this paper. I should also like to thank Dr P. Freeman, Dr J. F.
Perkins and Dr D. R. Ragge for their help and advice.

I am especially indebted to the following for the loan of type and other material
and the provision of valuable information in reply to my enquiries. Professor
R. H. Arnett, Jr., and Mr E. C. Mignot (Purdue University, Layfayette, Indiana),
Dr A. Capart and Dr R. Damoiseau (IRSNB, Brussels), Mr O. L. Cartwright,
Dr K. V. Krombein, Dr P. J. Spangler-and Dr T. J. Spilman (USNM, Washington),
Dr P. N. Chatterjee (FRI, Dehra Dun), Mr M. Clifton (MN, Nairobi), Colonel
C. F. Cowan (Little Gaddesden House, Berkhamsted, Herts), Professor P. F.
Darlington (MCZ, Harvard), Dr L. Diekmann and Dr R. Gaedike (DEI, Eberswalde),
Dr J. P. Duffels (ZM, Amsterdam), Professor S. Duncan (Boston University,
Mass.), Dr E. Esfandiari (Ministry of Agriculture, Tehran, Iran), Dr G. Hallin
and Dr T. Nyholm (NR, Stockholm), Dr F. Hieke (ZMHU, Berlin), D. H. Hippa
and Dr P. Lehtinen (TU, Finland), Dr J. O. Husing (ZIMLU, Halle-Wittenburg),
Dr F. Janczyk (NM, Vienna), Mr D. B. Janson (44, Great Russell Street, London),
Dr F. Keiser (NM, Basle), Dr Z. Kaszab (TM, Budapest), Dr S. Kelejnikova (ZMU,
Moscow), Dr O. Kryzhanovsky (ZI, Leningrad), Dr S. G. Larsson (UZM, Copen-
hagen), Mr H. B. Leech (CAS, San Francisco), Mr R. H. Lutts (Museum of Science,
Boston, Mass.), Dr M. Mroczkowski (IZPAN, Warsaw), Dr A. Neboiss (NMV,
Victoria), Dr H. Ohira (Aichi University, Okazaki City, Japan), Professor L. Pardi
and Professor U. Parenti (IMZU, Turin), Miss P. Schuyler (ANS, Philadelphia),
Dr H. Silfverberg (ZMU, Helsinki), Dr J. Stehik (MM, Brno), Mr E. Taylor (UM,
Oxford), Professor E. Tortonese and Dr D. Guiglia (MCSN, Genoa), Dr S. L. Tuxen
(UZM, Copenhagen), Madame Bons, Dr A. Villiers, Dr A. Descarpentries and
Dr C. Girard (MNHN, Paris), Dr G. E. Wallace (CM, Pittsburgh) and Dr J. Wiebes
(RNH, Amsterdam).

270 CaM. BF. von HAYEK
SOME NOTES ON THE COLLECTIONS CONSULTED

Where no reference is given the information concerning the present location
of a collection was obtained from Horn and Kahle (1935-1937) and confirmed
by correspondence with the present owner or by personal inspection of the collection.
References are given to all other sources of information.

Albertis, Luigi Maria d’ (1841-1901). See Beccari.

Alluaud, Charles (1861-1949). Madagascan material (except Carabidae) and
specimens collected in East Africa with R. Jeannel to MNHN, Paris.

Andrewes, Herbert Edward (1863-1951). Coleoptera (except Carabidae) to
BMNH.

Candéze (1893c) described a number of species from the H. E. Andrewes
collection. He appears to have retained the unique holotypes of some
species (e.g. Pevricus sanguinolentus) which are now in the IRSNB, Brussels and
returned half of longer series (e.g. 3 out of 6 L. pistoreus) to Andrewes.

Bakewell, Robert (1910-1867). Elateridae via Neervort van de Poll and
Fleutiaux to MNHN, Paris.

Bates, Henry Walter (1825-1892). Elateridae via Janson and Godman and
Salvin to BMNH.

Beccari, Odoardo (1843-1892). Material collected in Java, Celebes, Timor,
etc., with d’Albertis (1871-1876) to MCSN, Genoa. Candéze (1878a) described
a number of species from this material but up to the present time I have
been unable to discover whether he retained any of the specimens. If he
did, they should be in the IRSNB, Brussels. African collection to MCSN,
Genoa.

Blackburn, Thomas (1844-1912). Blackburn did not publish designations of
type-specimens or record the number of specimens on which he based his
descriptions. He was, however, in the habit of marking one specimen of
each species, with the letter ‘T’ in red ink. Up to the present time these
specimens have been accepted as types. In 1910 the BMNH purchased a
collection of 698 such ‘types’ from Blackburn. Since Blackburn obviously
intended his type-material to be preserved in the BMNH, I have designated
lectotypes from the material in that collection. The rest of Blackburn’s
collection, which presumably contains the remainder of any syntype series,
was acquired by the SAM, Adelaide and part by C. French (q.v.).

Van Zwaluwenburg (1959 : 352, under lindensis Blackburn) states that the
numbers on the card mounts of Blackburn’s specimens may perhaps refer
to a locality in Blackburn’s register deposited in the SAM, Adelaide. I have
not confirmed this.

Bosc d’Antic, L.A.G. (1759-1825). The Bosc collection was acquired by the

MNHN, Paris in 1828. Zimsen (1964: 17) believed that the Coleoptera had
been lost or destroyed. The discovery of the type-material of E. pennata

ee . a

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 271

Fabricius (p. 62) and £. varius Olivier (p. 79) has shown that this is not
the case.

Candéze, Ernest Charles Auguste (1827-1898). Candéze’s real interest lay in

the Lamellicorns, but Lacordaire persuaded him to undertake the classification
of the Elateridae. As he had no intention of continuing work on this family
once the Monograph was finished, Candéze did not build up a large collection
of his own but based his work on material sent to him on loan by Museums
and owners of private collections (1857: VII-VIII). Fortunately Candéze
made a habit of recording the names of the owners of the specimens he
described. In those cases where he does not mention the owner, I have
assumed that the material was in his own possession.

On completion of the Monograph in 1864 Candéze negotiated an exchange
with Janson, giving him some (‘quelques’, Fleutiaux, 1945 : 79) of the Elaterid
types in his possession in return for La Ferté Sénectére’s Lamellicorns. The
exchange took place in 1869. Janson (q.v.) appears to have retained Candéze’s
material which is now preserved in the BMNH. The BMNH possesses a
manuscript catalogue of the Janson collection made by Waterhouse. In it
Waterhouse records that ‘Mr Janson’s collection comprises La Ferte’s as well
as Candéze’s’.

The publication of the Monograph established Candéze as the acknowledged
Elaterid specialist and he found that he was unable to carry out his intention
of abandoning the family. Instead he embarked on a revision of the Mono-
graph, which unfortunately remained unfinished, and began to build up a
collection of Elateridae. At the time of his death he was the owner of a large
and comprehensive collection which included the Elaterid collections of
Lacordaire, Laporte de Castelnau, Reiche (8 boxes), Semper (Philippine
material), Buquet, Monchicourt (including Raffray’s Zanguebar collection),
Mniszech, Gebler, Falderman, Duport, Helmberg (Alaskan material), Wallace
(New Guinea, etc.) and Fairmaire’s exotic material. This second collection
which presumably included any specimens, including syntypes, retained in
1869, was purchased by the Belgian government in 1899 (1899, Annis Soc.
ent. Belge 34 : 31, report) and is preserved in the IRSNB, Brussels.

Candéze described a number of species from ‘des Indes-Orientales’.
Subsequent records of certain species (e.g. Adelocera tostus (= Agrypnus of
the present work) (Candéze), Fleutiaux, 1927 : 78) suggest that this rather
imprecise locality includes eastern India, Burma, Thailand, Indochina, Malaya
and Indonesia. In many cases Janson’s replacement locality labels bear
only the word ‘India’.

Fleutiaux (1945 : 81) states that Candéze was in the habit of retaining
the specimens submitted to him for identification. Experience (see note on
the Andrewes collection p. 270) has shown that this was not always the case.

Except in those cases in which he had only a single example, Candéze did
not generally record the number of specimens on which he based his description.
In addition he did not always record the exact locality on the specimen so
that the published locality and that on the type material do not always agree.

272

Cc. M. F. von HAYEK

Candéze was not consistent in the way in which he labelled his specimens.
In most cases he seems to have placed his determination label on the first
specimen of a series. Many Candéze specimens in the BMNH bear large
paper labels with coloured borders (blue = Africa, yellow = Asia, red =
Australia, etc.) with the generic and specific names together with those of
the author and collector written in his normal handwriting. On other
occasions he used a small scrap of paper on which he wrote only the specific
name. A large number of specimens in the IRSNB, Brussels, bear large
stiff card labels with coloured borders. One of these is illustrated in Horn,
1936: p. 21, fig. 22. The handwriting, a form of printing, differs from that
on the BMNH labels. I believe that these labels may have originally been
pinned in the boxes above or below the species rather than attached to the
specimens themselves. Candéze sometimes wrote the word ‘type’ on his
determination labels, but he does not appear to have done so for every species
he described.

It is unfortunate that when Janson acquired part of the Candéze collection
he considered it necessary to replace Candéze’s determination labels with his
own. Sometimes Candéze’s labels have been trimmed and stuck to the
underside of the new labels, but more often they seem to have been removed.

Fleutiaux (1945) published a detailed account of the history of the Candéze
collection.

Cantor, Dr Theodore Edward (1809-1860). Dr Cantor of the Bengal Medical

Service accompanied the military expedition to Chusan [Chushan, China]
with instructions to collect objects of Natural History for the Court of Directors
of the East-India Company (Cantor, 1842 : 368). The expedition travelled by
way of Penang, Singapore and the island of Lantao in the Canton River,
where it remained for the whole of the month of June, 1840. The occupation
of Chusan lasted from July 1840 to March 1841. A collection of insects made
in Chusan was presented to the East-India Company in 1842 (Cantor, 1842 : 278
and Horsfield and Moore, 1857: 2) and a duplicate series was sent to the
Entomological Society of London (Cantor, 1842 : 278).

The BMNH acquired part of the East-India Company’s collection when it
was broken up in 1860. The BMNH also acquired the type-material belonging
to the Entomological Society before its collections were sold by auction
between 1859 and 1863.

Hope (1843) described a number of species from Chusan and Canton collected
by Cantor but does not record whether the specimens belong to the East-India
Company, the Entomological Society or whether he had acquired them himself.
Hope seems to have had some of Cantor’s material in his possession as two
specimens believed to be the types of two species (Melolontha chinensis and
Anomala controversa) described by Hope in 1843 are in the UM, Oxford
However Candéze’s comments (1859 : 235, 405 and 1863: 242) that he saw
the types of Ludius luteipes, crocopus and quadrilineatus Hope, 1843 in the
Indian Museum and Crotch’s (1873 : 41) remark that he inspected the types

?

b
>

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 273

of Coccinella 18-spilota, succinea and tetraspilota Hope, 1843 in the same
museum leaves little doubt that Hope probably based most of his descriptions
on specimens belonging to the East-India Company.

Inspection of the BMNH collection has shown that it contains specimens
regarded by a number of different workers as the types of 39 of the 49 species
described by Hope in 1843. A few specimens lack registration labels but the
remainder bear labels indicating that they were acquired from the East-India
Company. Most specimens bear a small round label with the word ‘China’ on
one side and ‘Dr Cantor’ on the reverse but a few are labelled ‘Penang’.

The unknown author of a short biography of Cantor cut from an unidentified
publication and pasted on three cards in the BMNH library card index remarks
that ‘many of his [Cantor’s] data as regards the locality—particularly those
labelled Penang are incorrect’. The writer appears to be referring to Cantor’s
collection of reptiles and amphibians but I suspect (see orientalis Hope p. 192)
that the same may apply to the insect collection.

Cardon, René P. (1879-[date of death unknown). Pére Cardon, Jesuit missionary
in India and keen insect collector, sent specimens collected in Chota Nagpore,
Tetara, Konbir-Nawatoli and later at Barwai to his friend and colleague Pére
Renard, Préfet des Etudes du Collége St Servan at Liége (Candéze, 1890:
CXLVII). These collections have not been located, but it seems probable that
Candéze acquired the Elateridae. The IRSNB, Brussels, Oberthur, Fleutiaux
and Fry collections all contain specimens collected by Cardon and determined
by Candéze.

Specimens believed to form part of the syntype series of certain species
(e.g. succinatus Candéze, p. 48) do not bear locality labels corresponding
to the published type locality. Instead they are labelled ‘Bengal’ or ‘Chota
Nagpore’. It is assumed that Candéze obtained more detailed information
from Cardon.

Candéze (1890 : 46) describes Chota Nagpore as ‘un province du Bengale
occidentale s’entendent au sud de la grande plaine du Gange’ and two years
later (1892c : 481) he quotes Cardon’s companion P. Maene, S.J., who describes
their parish as ‘Le Barwai’ [=Barwa 23°12'N., 84°18’E.], le Chechari, le Kasir
et une partie de Noaghar . . . qui sont compris entre 22°58’N. and 33°30’N.
and 84°E. and 84°30’E.’ The area is traversed by the rivers Sunk [=Sankh]
and Koel.

Castelnau, Francis L. N. de C. de Laporte. Comte de (1810-1880). Material
from North and South America to MNHN, Paris. First private collection
to NMV, Melbourne. The Elateridae of his second collection, which were
in rather poor condition, were offered for sale by Deyrolle and purchased
by Candéze (q.v.) in 1872.

Some writers refer to this worker as Laporte. Musgrave (1932 : 180) states
that at first he wrote under the name of Laporte or Delaporte but later that
of Castelnau. Both the Schenkling catalogue (1925) and the Derksen and

274 CoM BE Won’ ELAWAERIS

Scheiding bibliography (1963) use the name Castelnau. Personal preference
is the reason for the use of Castelnau is this work.

Chevrolat, Louis Alexandre August (1799-1884). At his death his Elateridae
were acquired by Fleutiaux and are now in the MNHN, Paris. Fleutiaux
retained Chevrolat’s large coloured determination labels. The specimens
also bear a printed ‘Collection Chevrolat’ label.

David, Armand (1826-1900). His first collection was destroyed. Later col-
lections were broken up: part was acquired by the MNHN, Paris, part by
Oberthur (now in MNHN, Paris) and part by the Institut Catholique de Paris
[not confirmed]. At his death his private collection was acquired by Oberthiir.

Dejean, Pierre Francois Marie Auguste, Comte (1780-1845). Elateridae via
H. L. Gory, La Ferté Sénectére, Janson and Godman & Salvin to BMNH.

Deyrolle, Achille (1813-1865). Elateridae passed via Janson to BMNH. I have
assumed that the Deyrolle to whom Candéze refers in the Monograph is
A. Deyrolle and not his son Emile or brother Henri.

Dohrn, Carl August (1806-1892). His collection was acquired, part direct and
part through his son, by the Museum fiir Naturkunde, Stettin (Szcezin). At
least part of the Stettin collection is now preserved in the IZPAN, Warsaw.

Dupont, E. ( ) and (Richard) Henry (1798-1873). The Dupont brothers
were dealers in natural history specimens trading in Paris. They also under-
took collecting expeditions. Germar (see Agrypnus terrenus (=crenatus Klug)
p- 144, turbidus, p. 224, probably obtained some of his material from the
Duponts. In 1846 Mniszech (q.v.) obtained a collection of Elateridae from
Dupont (Candéze, 18950 : 51) and another collection acquired by Reiche passed
to Janson (q.v.).

Elston, Albert H. (1890- ). Elston generally based his descriptions on an
unrecorded number of specimens from several localities adding ‘type in the
author’s collection’ or ‘in the South Australian Museum’ [Adelaide]. He did
not record the location of the rest of the material. Elston’s collection is
now in the AM, Sydney. Neboiss’s (1956, 1961) records suggest that where
Elston’s ‘type’ is in his own collection, some or all of the remaining specimens
were acquired by the SAM, Adelaide, and where the ‘type’ was in that museum,
some or all of the remaining material was retained by Elston and is now in
the AM, Sydney.

Since Elston did not publish any means of identifying his types, lectotype
designations should be made, but with due regard to Elston’s intention con-
cerning the location of the type. In this work, where Elston’s specimens
are preserved in more than one place the first named corresponds to the
location of Elston’s ‘type’.

Eschscholtz, Johan Friedrich (1793-1831). Collection in ZMU, Moscow and
also some material in ZMU, Helsinki. Not confirmed.

~ st hae

.
;

|
|
|
|

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 275

Fairmaire, Léon (1820-1906). Fairmaire appears to have disposed of parts of

his collection during his lifetime. Candéze acquired a collection of exotic
Elateridae in 1894 (now in the IRSNB, Brussels) and Fleutiaux acquired a
second collection of exotic and American specimens (now in the MNHN, Paris).
The collection in his possession at the time of his death is in the MNHN, Paris.

Fea, Leonardo (1852-1903). Explorer and naturalist. The main part of his

collection is in the MCSN, Genoa. Duplicates were acquired by other museums
and private collectors. An account of his travels in Burma will be found
in Fea (1896). Candéze (1891 : 771) records a number of species from Carin
Cheba collected by Fea. He describes the locality as ‘Les montagnes des
Carin, a une cinquantaine des lieues au nord du Rangoon et a une hauteur
de 900 a 1500 metres.’ This is presumably the district of Burma now known
as Karen.

Fleutiaux, Edmond (1858-1951). At his death his large and valuable collection

was acquired by the MNHN, Paris. It contains specimens from many other
collections. In most cases these are easily recognizable, as Fleutiaux retained
all original labels and often added his own to indicate the provenance of a
specimen. The collection also contains notes on type-material in other collec-
tions. Fleutiaux did not generally record the number of specimens on which
he based his description, nor did he designate types. However, he was in
the habit of marking one specimen as the type, usually by writing the word
‘type’ on the determination label. Wherever possible I have designated the
specimen so marked as the lectotype.

French, Charles (1840-1933). First collection via Lansberge and Oberthiir to

the MNHN, Paris. Second collection to Neervoort van de Poll (q.v.). Third
collection to NMV, Melbourne.

Germain, Philibert (1827-1913). Elateridae via Janson and Godman and Salvin

to the BMNH. Some of his material was acquired by Oberthiir (q.v.) and
the Museo Nacional, Santiago, Chile (not confirmed).

Germar, Ernest Friedrich (1786-1853). Germar’s collection was broken up.

The Curculionidae passed to the University Zoological Museum, Halle (now
the Zoological Institute of the Martin Luther University, Halle Wittenburg).
This is confirmed by Dr Husing, who adds that the collection does not contain
any Elateridae. The remainder of the collection passed to his nephew, Dr H.
Schaum. Schaum seems to have retained some of the Elateridae (see p. 280)
but he appears to have disposed of material belonging to other families (see
Horn & Kahle, 1936: 241) partly to the NMHU, Berlin and partly to the
DEI, Eberswalde.

Gerstaecker, Carl Edward (1828-1895). Curator at the MNHU, Berlin, until

about 1876. He appears to have relabelled the Coleoptera collection, replacing
many old labels including those of Klug, Herbst and Germar.

276 Cz Mo PY Von SHAY EK

Goudot, Justin ( ). Dealer in natural history specimens in Paris.
Lived and collected for some time in Tananarive, Madagascar. In 1830
(Candéze, 18950: 50) he sent specimens from Imerina to H. Dupont (q.v.),
the MNHN, Paris and MNHU, Berlin.

Harris, Thaddeus William (1875-1856). Say based the description of several
species (e.g. Elater obtectus, aurorata) on specimens submitted by Dr Harris.
The Harris collection passed to the Boston Society of Natural History. Dr R.
Lutts, curator of collections at the Boston Museum of Science informs me
that in the late 1940’s the Society’s collections were transfered to a number
of other institutions. Professor P. J. Darlington has informed me that the
Harris collection, and also his notebooks, are now in the MCZ, Harvard. The
collection appears to contain only one Agrypnine specimen which could be
part of Say’s type-material. (See Lacon auroratus (Say) p. 57).

Horn, George Henry (1840-1879). Coleoptera to the ANS, Philadelphia.

Janson, Edward Wesley (1822-1891). Dealer in natural history specimens
in London. His collection of Elateridae which included the Elateridae from
many older collections was acquired by Dr F. D. Godman who presented it
to the BMNH. The specimens bear the registration number 1903.130. Janson
is known to have been in the habit of replacing old labels with his own. On
these he generally recorded the generic, specific and author’s names, and also
the name of the person responsible for naming the specimen. He often included
the name of the original owner of the specimen, e.g. Lacon occidentalis Cdze.
Cand. [identified by] Type e coll. de Laferté [original owner].

Kraatz, Gustav (1831-1909). Collection acquired by the DEI, Eberswalde.

Knoch, August Wilhelm (1742: 1818). Zincken (1818) published a short biography
in which he records that Knoch’s collection, which appears to have been
quite a large one, was for sale at 1,000 Thaler in gold. Hagen (1862 : 425)
records that the collection is in the ZMHU, Berlin, but Dr Hieke writes that
the Museum appears to possess only part of the Knoch collection. Unfortu-
nately the records of how the collection was acquired and its contents cannot
be found. What happened to the remainder of the collection is unknown.

La Ferté—Sénectére, F. Thibault de (1808-1866). Part of his collection was
acquired by Janson in 1864. Janson sold most of the families separately
but retained the Elateridae which passed, with his collection, to the BMNH.
Some specimens bear a BMNH registration label ‘Janson Coll. ex. La Ferté
1903.130’ while others have the ordinary Janson coll. registration labels with
Janson’s replacement locality labels with the abbreviation ‘Laf.’ or ‘e coll.
Laferté’.

Lansberge, Johan Wilhelm van (1830-1905). Coleoptera to Oberthiir (q.v.).
Duplicate material to IRSNB, Brussels, and Neervoort van de Poll (q.v.).

Laporte, Francis L. de, Comte de Castelnau, see Castelnau.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 277

Latreille, Pierre Andre (1762-1833). His first collection was acquired by Dejean
(q.v.) in 1826. The Elateridae of the second collection were acquired by
Janson (q.v.).

LeConte, Henry (1825-1883). North American Coleoptera to MCZ, Harvard.
LeConte’s specimens bear small coloured paper disks. Professor P. J.
Darlington informs me that the colours are believed to indicate the following

localities:

pink = Middle States

brick red or orange = Southern States

yellow = Western States

dark red = Devas

light green = Nebraska & C [so forth]. In the past Nebraska was
the name of a much larger area.

dark green = New Mexico

blue = Oregon & C. Oregon territory formerly included Wash-

ington State
light blue-grey = Lake Superior

: Lewis, George (1839-1926). Lewis’s collection of Coleoptera from Japan, China
and Ceylon was acquired by the BMNH in four instalments which were
registered as follows: IgI0 : 320, Ig1I : 298, 1912 : 276, 1913 : 256.
: Lewis paid two visits to Japan, the first in 1867-1872 and the second in
1880-1881. Bates (1883) gives an account and detailed itinerary of the second
visit.
Lewis submitted the Elateridae collected in the course of the first visit to
Candéze, who published a paper based on this material in 1873. Lewis (1894:
36) records that Candéze returned the specimens to him but the presence of
| specimens from Lewis’s first journey in the IRSNB, Brussels, suggests that
Candéze retained representatives of some species. Some of the specimens
may be syntypes. Lewis appears to have relabelled some of the specimens
returned by Candéze. For example, Lacon twmens Candéze (see p. 49) was
described from a single specimen. The BMNH possesses a single male from
the Lewis collection which agrees well with the description but bears a determin-
ation label., ‘Lacon tumens n. sp. Cdze’ in Lewis’s handwriting. Similarly
the specimen of L. scrofa (see p. 208) which most closely resembles the descrip-
tion is in the Lewis collection without Candéze’s determination label but with
Lewis’s label ‘Lacon scrofa n. sp. Cdze’.
Lewis described the Elateridae collected on the second expedition himself.
Except in those cases in which he possessed only a single specimen, he did not
generally record the number of specimens he had before him at the time of the
description. His remark (1894: 26) that ‘the localities given are selected
to furnish some information regarding the distribution of the species’ suggests
that he did not record the localities of all the specimens on which the description

278

CoM sB. von, HAW EK

was based. Lewis did not place a determination or locality label on every
specimen of his series nor was he consistent in marking one specimen as the
type. Where there is sufficient evidence that a particular specimen formed
part of Lewis’s original series this specimen has been treated as a syntype
even though the locality on the label differs from that published or there is
no locality label at all.

Long, Major S. H. For an account of the expedition from Pittsburgh to the

Rocky Mountains see James (1823). The material collected on this expedition
has not been traced. See also Say, p. 279.

Mniszech, George Vandelin (1824-1881). Mniszech’s collection was purchased

by Lansberge, who sold the Elateridae to Candéze (q.v.) in 1871 (Candéze,
1895 : 51, Fleutiaux, 1945 : 80). This material is now in the IRSNB, Brussels.

Mniszech acquired some of Dupont’s Madagascan material in 1846. His
collection also contained those of Gebler, Falderman, Helmberg, Kindermann,
Deyrolle and many specimens collected by Wallace in New Guinea. According
to Fleutiaux (1945 : 81) all the old labels were removed for display purposes.

Candéze described a number of species from Mniszech’s collection before
it came into his possession. It would appear that in some cases he retained
the specimens (e.g. Lacon nodicollis, see p. 191, L. sinuatus, see p. 213) or
part of the series (e.g. Lacon eximus see p. 152) on which he based the desc1ip-
tions as they were acquired by the BMNH with his first collection and are not
in his second collection in the IRSNB, Brussels.

Modigliani, Elio (1860-1932). Collections made in Nias, Sumatra, Enggano

(May—June 1891) and Mentawei are in the MCSN, Genoa. Any specimens
described and retained by Candéze should be in the IRSNB, Brussels.

Mochul’skii, V. I. de. See Motschulsky.

Motschulsky, Victor Ivanovic de (c. 1810-1871). The Russian Coleopterist

used this form of his surname in both the publications consulted in the course
of this work. For the last hundred years or more coleopterists have used
this spelling and the abbreviation ‘Motsch’. In my opinion no useful purpose
is served by changing the familiar spelling.

Horn (1936 : 183) states that the Motschulsky collection is in the ZMU,
Moscow, and the Zoological Museum, Leningrad, while the Naturforschender
Verein, Riga, the MNHU, Berlin and the DEI, Eberswalde, possess ‘duplicates’.
However enquiries have shown that neither of the German institutions possess
Coleoptera determined by Motschulsky.

Fleutiaux (1932e: 79) records that the Motschulsky collection suffered
considerable damage before it was acquired by the ZMU, Moscow. He does
not mention any material in other institutions. Fleutiaux was fortunate
in that through the kindness of the curator, Mr Kuzin, he was able to examine
the specimens regarded as Motschulsky types and his collection contains a
number of specimens of Motschulsky species labelled ‘comparé au type’.

:
;

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 279

Motschulsky did not record the number of specimens on which he based his
descriptions nor did he designate type-specimens.

Murray, Andrew (1812-1878). Received collections of beetles made by the
staff of the Presbyterian missionary station at Old Calabar [Nigeria]. He
submitted (1857: 153) many Elateridae to Candéze. Whether Candéze
returned any or all of the material to Murray is unknown. On his death,
Murray’s collection was sold by auction. Janson (q.v.) acquired the Elateridae.
Any specimens retained by Candéze should be in the IRSNB, Brussels.

Neervort van de Poll, Jacob R. H. (1862-1925). Exotic Elateridae via Fleutiaux
to the MNHN, Paris.

Oberthiir, René (1852-1944). The Oberthiir collection was declared a National
Monument in 1952 and is now housed in the MNHN, Paris. A note in the
Entomologische Bldtter 55 (1960) : 282 states that part of the Oberthiir collec-
tion, including a number of Coleoptera, was acquired by the Museum A. Koenig
in Bonn. Dr H. Roer informs me that this material consists of a collection
purchased from Oberthiir by Dr Hone. The Agrypninae in the collection
lack locality labels.

Oberthiir retained only the best specimens of the collection he purchased,
re-selling the rest to Neervort van de Poll (q.v.) and H. Donckier, a dealer
in insects. Oberthiir does not appear to have been interested in the Elateridae.
His collection includes some 150 boxes containing both determined and undeter-
mined Elateridae from all parts of the world. As they are found, type-material
and determined specimens are being transferred, with labels to show their
provenance, to the general collection of the MNHN, Paris.

Philippi, Rudolph Amandus and Friedrich Heinrich Euonom (1838-1910, father
and son). Collection to the Museum Nacional, Santiago, Chile (not confirmed).
R. A. Philippi was at one time director of the Museum Nacional, Santiago.

Raffray, Achille (1844-1923). Coleoptera of Abyssinia and Ethiopia via F.
Monchicourt and E. Candéze to the IRSNB, Brussels.

Sallé, Auguste (1820-1896). Dealer in natural history objects in Paris. As
a boy (about 1831-34) he collected, probably for Chevrolat, in the Southern
U.S.A., W. Indies, Central America and Venezuela. His own collection of
Central American and Mexican Elateridae was acquired by Godman and
Salvin and presented to the BMNH.

Saunders, William Wilson (1809-1879). His collection was broken up by Janson,
who retained some of the Elateridae (e.g. Lacon comptus Candéze, see p. 142).
The remainder was acquired by Neervort van de Poll (q.v.).

Say, Thomas (1887-1834). Both LeConte (1859a: VI) and Ord (18594: XIX,
footnote) record that Say’s collection was entirely destroyed. Horn (1936 : 240)

records that such as remains of Say’s collection is in the ANS, Philadelphia,

280 CaaS Es WON ee Aw Ek

but Miss P. Schuyler reports that the collection does not contain any beetles
from the Say collection.

Say described several species based on specimens he collected while he was
with Major Long’s expedition to the Rocky Mountains in 1819~20 (Say, 1825 :
259). It seems probable that he retained them and that they were destroyed
with his collection. However, if he did not retain them, they may have passed,
with other material collected on this expedition, to Peales Philadelphia Museum
(Horn & Kahle 1937 : 369). This collection suffered many vicissitudes and
was finally broken up (Colston, 1909). Part passed, via the Boston Museum
to the Boston Society of Natural History and part to Barnum’s American
Museum which was destroyed by fire. Mr R. Lutts, Curator of collections at
the Museum of Science, Boston, where the Boston Society collection was
housed, has informed me that in the early 1940’s the bulk of the Society’s
research material was transferred to other institutions including Boston and
Harvard Universities. Mr Lutts adds that he can find no reference to Major
Long’s expedition or Peale’s Philadelphia Museum material in the accession
records. There seems little hope of tracing this material.

Schaum, Herman Rudolph (1819-1865). A nephew of E. F. Germar, he was
also interested in the Elateridae and appears to have retained at least part
(see Candéze, 1857 : 132 and 100) of his uncle’s Elaterid collection. Part of
Schaum’s collection was acquired by Janson and is now in the BMNH. What
became of the rest of the Schaum collection is unknown. The BMNH possesses
a manuscript list entitled ‘Catalogus Elateridarum collectionis Schaumii’.
It appears to be a list of the Schaum Elateridae acquired by Janson. Only
a small proportion of Germar’s species are included in the list.

Schwarz, Otto (1861-1908). Elateridae via O. Leonhard to the DEI, Eberswalde.
Schwarz did not publish type-designations nor did he record the number of
specimens on which he based his description or where they were preserved.
Unless there is evidence to the contrary, I have assumed that his type-material
is in his own collection.

Ulke, Henry (1821-1910). Coleoptera to CM, Pittsburgh.

Wahlberg, Johan August (1810-1856). Collected in ‘Caffraria’ from 1838 to
1845. For an account of his itinerary see Brink (1935 : 25-34). The material
is now in the N.R., Stockholm. Boheman described Lacon amplicollis and
troglodytes from ‘tractitus fluvii Gariepis’ [Orange R.], nanus, paenulatus and
decipiens from ‘Terra Natalensi’ and parcus from ‘regione fluvii Limpopoensis’.
Through the kindness of Dr Hallin, I have been able to examine all the syntype
material of these species. None of the specimens bear labels corresponding
to the published locality, only ‘Caffraria’ and ‘J. Wahlb.’. In each case the
specimen with Boheman’s determination label bears a type label which I
am given to understand was added at a later date as a matter of curatorial
routine.

From Brink’s account of Wahlberg’s travels it would appear that the term

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 281

‘Caffraria’ is sufficiently wide to include all three localities. Pope (1960 : 206,
210) is of the same opinion.

Xantus de Vesey, John ( ). Horn (1894 : 302) records that the material
collected in the Californian peninsula in 1859 and 1860 was deposited in the
Smithsonian Institution and afterwards divided between Ulke (q.v.) and
LeConte (q.v.).

The locality of the Xantus material is usually described as Cape St. Lucas.
According to Horn (loc. cit.) this material was obtained in the region between
San José del Cabo and La Paz.

REFERENCES

Where an author has published more than one work in the same year these have been listed
chronologically. Where only the year of publication could be discovered the date of publication
is interpreted as the last day of December of that year.

ARNETT, R. H. 1952. A review of the Nearctic Adelocerina (Coleoptera, Elateridae, Pyro-
phorinae, Pyrophorini). Wasmann J. Biol. 10 : 103-126.

1953. Notes on the genus Lacon (Elateridae). Coleopts Bull. 7 : 5-7.

1955. Supplement and corrections to J. A. Hyslop’s Genotypes of the Elaterid beetles
of the world. Pyvoc. U.S. natn. Mus. 103 : 599-619.

ARNETT, R. H., Micnot, E. C. & Smitu, E. H. 1969. North American Coleoptera fauna;
notes on Pyrophorinae, Elateridae. Coleopts Bull. 23 : 9-15.

Asso, I. pg, 1801. Introduccion 4 la Ichthyologia oriental de Espatia. An. Cienc. nat. Madrid
4: 28-52.

Basaurt, G. 10917. Chasses et recherches zoologiques en Afrique orientale anglaise 1973.
213 pp. Illustrations and map. Paris.

Bates, H.W. 1866. Ona collection of Coleoptera from Formosa, sent home by R. Swinhoe,

Esq., H. B. M. Consul, Formosa. Proc. zool. Soc. Lond. 1866 : 339-355.

1883. Supplement to the Geodephagous Coleoptera of Japan, chiefly from the collection
of Mr George Lewis, made during his second visit from February, 1880, to September, 1881.
Trans. ent. Soc. Lond. 16 : 205-290, 1 map, 1 pl. An account of Mr Lewis’ itinerary is
given on pp. 210-213.

BavupDI Di SELVE, F. 1871. Coleopterorum messis in insulo Cypro et Asia minore ab Eugenio
Truqui congregatae recensitio. Berl. ent. Z. 15 : 40-71.

BeEQuaeERT, J. 1926. The date of publication of the Hymenoptera and Diptera described by
Guérin in Duperrey’s ‘Voyage de la Coquille’. Ent. Mitt. 15 : 186-195.

BinaGuI, G. 1941a. Gli apparati genitali dell’ Agrypnus notodonta Latr. e descrizione di

una nuova specie di Agvypnus dell’ Africa settentrionale. Boll. Soc. ent. ital. 73 : 68-74,

10 figs.

1941b. Revision dei Tipi di E. Candéze delle collezioni del Museo di Genova. Nota |.
Boll. Soc. ent. ital. 73 : 83-89.

1941c. Revisione dei Tipi di E. Candéze delle collezioni del Museo di Genova. Nota II.
Boll. Soc. ent. ital. 73 : 118-123.

BLACKBURN, T. 1890. Further notes on Australian Coleoptera with descriptions of new
genera species [sic]. Tvans. R. Soc. S. Aust. 13 : 82-83.

—— 1892a. Notes on Australian Coleoptera with descriptions of new species. Proc. Linn.
Soc. N.S.W. (2) 6 (1891) : 479-550.

1892b. Notes on Australian Coleoptera with descriptions of new species. Proc. Linn.

Soc. N.S.W. (2) 7 : 283-300.

1895. Further notes on Australian Coleoptera with descriptions of new genera and
species. Tvans R. Soc. S. Aust. 19 : 27-60.

282 CLM BS “won Gl ANGE

BLACKBURN, T. 1900. Further notes on Australian Coleoptera with descriptions of new genera
and species. Tvans R. Soc. S. Aust. 24 : 35-68.

BLANCHARD, E. 1845. Jn d’Orbigny, Voyage dans l Amérique meridionale. 6 part 2.
Insects de l Amérique méridionale recueillis pay A. d’Orbigny. 222 pp. [1836-47], 32 pls
[1835-1845]. M. Brullé was the author of pp. 1-57 (see introductory note). The dates of
publication of the separate parts were obtained from Sherborn & Griffin (1934).

1853. Jn Dumont d’Urville, Voyage au Péle Sud et dans lOcéanie sur les Corvettes
U Astrolabe et la Zélée. Zoologie 4. 422 pp. Atlas, 140 pls (1842-53). The year of the
publication of the individual plates is unknown to me.

BLACKWELDER, R. E. 10944. Checklist of the Coleopterous insects of wees Central
America, the West Indies and South America. Part 2. Bull. U.S. natn. Mus. 185 : 180—
Sac

BLatcHLEy, W.S. i910. The Coleoptera of Indiana. 1386 pp., 590 figs. Indianapolis.

BowHEMAN, C. 1851. Insecta Caffraria annis 1838-45 a J. A. Wahlberg collecta . . . Coleoptera
1, fasc. 2, 299-626; 2 pls. Stockholm.

Botspuval, J.B. A.D.bDE. 1832. Voyagede...l Astrolabe... pendant ...1826-29. Faune

entomologique de l’Océan Pacifique. 1. vii + 267 pp. Paris.

1835. Op. cit. 2. vii + 716 pp.

Brancsik, K. 1893. Beitrage zur Kenntnis Nossibé’s und dessen Fauna nach Sendungen
und Mittheilungen des Hern P. Frey. Jh. naturw. Ver. Trencsiney Kom. 15-16 (1892—
1893) : 202-258, 4 pls.

Brinck, P. 1955. Swedish exploration of South African animal life during 200 years. S.
Afr. anim, Life. 1 : 11-62.

BuURMEISTER, H. 1855. Handbuch dey entomologie. 4, part 2., 569 pp. Berlin.

Buysson, H. pu. 1891. Espéces nouvelles d’Elatérides. Echange 7 : 133-136.

1893. Faune Gallo-Rhénane, 496 pp. Caen. This work was published in parts with
separate pagination, as supplements to the Revue d’Entomologie between October, 1892
and March, 1906.

1911. Matériaux pour servir A la faune des Elatérides de la Haute et Basse Egypte.
Bull. Soc. ent. Egypte 2 : 17-47.

1920. Contribution a la faune du Maroc et du Nord de 1’Afrique (2@€me Note). Bull.
Soc. Hist. nat. Afr. N. 11 : 12-15.

CALLAN, E. McC. 1964. Ecology of sand dunes with special reference to the insect com-
munities. In Monographiae Biologicae 14. Ecological studies in Southern Africa, pp. 174-
183. Ed. D. H.S. Davis and others. The Hague.

CALVERT, G. BarTLETT. 1889. Monografia de los Elatéridos de Chile. An. Univ. Chile.
[Known to me only as a separate of 84 pp.] Santiago, 1808.

CANDEZE, E. 1857 (Mai). Monographie des Elatérides 1. viii + 400 pp., 7 pls. Mém.
Soc. v. Sci. Liége 12.

-~—— 1863. Monographie des Elatérides. 4. 534 pp.,6 pls. Mém. Soc. r. Sci. Liége 17.
1865. Elatérides Nouveaux 1. Mém. cour. Acad. r. Sci. Belg. (in 8°), 17, no. 1. 63 pp.
1873. Insectes recueillis au Japan par Mr G. Lewis. Elatérides. Mém. Soc. r. Sci. Liége
(2) 5, memoire no. 2. ii + 32 pp.

1874. Revision de la monographie des Flatérides. Mém. Soc. v. Sci. Liége (2) 4, mémoire
no. I. vili + 218 pp.

1875. Relevé des Elatérides des Iles Philippines avec les diagnoses de quelques espéces
inédites. Annls Soc. ent. Belg. (Bulletin) 18 : CXKVIII-CXXXV.

1878a (March). Relevé de BlatGedes recueillis dans les Iles Malaises, 4 la Nouvelle-
Guinée et au Cap York, par MM. G. Doria, O. Beccari et L. M. D’Albertis. Annali Mus.
civ. Stor. nat. Giacomo Doria 12 : 99-143.

18786 (month unknown). Elatérides Nouveaux 2. Annis Soc. ent. Belg. (Bull.) 21:
LI-CXLIII. Also issued as a separate, pp. 554, cover and title page. The reference to
the page of the separate is given in parentheses after the main reference.

1880. Addition au relevé des Elatérides Malais. Elatérides recueillis dans les régions

A

tne

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 283

avoisinant le détroit de Torres, par M. L. M. d’Albertis. Annali Mus. civ. Stor. nat.
Giacomo Doria 15 : 188-108.

1882. Elatérides Nouveaux 3. Mem. Soc. r. Sci. Liége (2) 9. ii + 117 pp.

1883. Deux Elatérides Nouveaux des Iles de la Sonde. Notes Leyden Mus. 5 : 204-206.

1887a (April). Quatre espéces nouvelles d’Elatérides de Ile de Sumatre. Notes Leyden
Mus. 9 : 189-101.

18876 (October). Description d’Elatérides nouveaux, provenant de Normantown: Baie
de Carpentaria (Australie septentrionale). Notes Leyden Mus. 9 : 285-290.

1888. Elatérides recueillis en Birmanie et au Tenasserim par M. L. Fea pendant les
années 1885-1887. Annali Mus. civ. Stor. nat. Giacomo Doria (2) 6 : 607-689. Also
issued as a separate of 23 pp. The reference to the page of the separate is given in
parentheses after the main reference.

1889. TElatérides nouveaux 4. Anmnls Soc. ent. Belg. 33 : 67-123. The references given
in the Schenkling catalogue suggest that this paper was also issued with separate pagina-
tion. The separate is unknown to me. The page numbers given by Schenkling will be
found, in parentheses, following the main reference.

1890. Note sur les Elatérides du Chota—Nagpore. Annis Soc. ent. Belg. (Bull.) 34:
CXLVI-CLVII.

1891a (March). Elatérides recueillis en Birmanie en 1888 par M. L. Fea. Annali Mus. civ.
Stor. nat. Giacomo Doria (2) 10 : 771-793.
1891b (October). Description de neuf Elatérides nouveaux du Musée de Leyde. Notes
Leyden Mus. 13 : 243-247.

1891¢ (month unknown). Catalogue méthodique de Elatévides connus en 1890. xii +
246 pp. Liége. *

1892a (September). Elatérides recueillis par M. Modigliani dan l’Ile d’Engano, en Mai
et Juin 1891. Annali Mus. civ. Stor. nat. Giacomo Doria (2) 12 : 795-800.

1892b (October). Viaggio de L. Loria nella Papuasia Orientale VI. Flatérides recueillis
par M. Loria en 1889-1891 dans la Nouvelle-Guinée méridionale et régions voisines.
Annali Mus. civ. Stor. nat. Giacomo Doria (2) 12 : 801-805.
1892c. (month unknown). Deuxiéme note sur les Elatérides du Chota-Nagpore. Anmnls
Soc. ent. Belg. 36 : 480-495.

1893a (February). Cinq espéces nouvelles d’Elatérides de l’Ile de Java. Notes Leyden
Mus. 15 : 125-127.

1893) (month unknown). Flatérides nouveaux 5. Mem. Soc. r. Sci. Liége (2) 18. 4éme
memoire. 75 pp.

1893c (month unknown). Additions aux Elatérides des Indes Orientales. Annis Soc.
ent. Belg. 37 : 168-179.

18954 (Feb.). Elatéridesde Togo. Dt. ent. Z. : 47-48.

1895) (month unknown). Mission Scientifique de M. Ch. Alluaud dans le Territoire de
Diego Suarez (Avril-Aoait 1893). Les Elatérides de Madagascar. Annis Soc. ent. Belg.
39 : 50-69.

1895¢ (month unknown). Elatérides recueillis par M. le docteur E. Modigliani aux bords
du lac de Toba, 4 Sumatra, de Novembre 1890 a Mars 1891. Annali Mus. Civ. Stor. nat.
Giacomo Doria (1894) (2) 14 : 484-504.

1897. Elatérides Nouveaux 6. Mem. Soc. r. Sci. Liege (2) 29, z2eme memoire. 88 pp.

1900 [posthumous]. Elatérides Nouveaux 7. Annis Soc. ent. Belg. 44:77-101. The
references in the Schenkling catalogue (1925) suggest that this paper was also issued as a
separate with pages numbered 1->. It is unknown to me. Schenkling’s page numbers
are given in parentheses after the main reference.

Cantor, T. 1842. General features of Chusan with remarks on the flora and fauna of that

| Island. Ann. Mag. nat. Hist. 9 : 265-278, 361-370, 481-4093.

| CarTER, H. J. »1939. Australian Coleoptera. Notes and new species. Proc. Linn. Soc.
N.S.W. 64 : 279-230, 2 pls.

284 G..M. F. von HAYEK

Carus, J. V. & ENGLEMANN, W. 1861. Bibliotheca zoologica. Zool. 2144 pp. Leipzig.
CasTELNAU, F. L. DE LaporTE, CoNTE DE. 1836. Etudes entomologiques. Revue Ent.
(Silbermann) 4 : 6-60, table.

1840. Histoive naturelle des insectes, Coléoptéres. 1. cxxv + 324 pp., 155 pls. Paris.
CHAMPION, G. C. 1894. Biologia cent.-am. Insecta, Coleoptera. 3 part 1. Elateridae:
258-551, 12 pls (nos 12-23). London.

1917. Notes on the Coleoptera etc. recorded from ‘Resin Animé’ by the Rev. F. W. Hope.

Entomologist’s mon. Mag. (3) 3 : 244-246.

CHEREPANOV, A. I. 1957. See TSCHEREPANOV.

CHEvroLaT, L. A. M. 1852. NRappel des Coléopterés décrits par Palisot de Beauvois aux

genre actuellement adoptés avec synonomie. Avnnls Soc. ent. Fr. (2) 10 : 629-651.

1867. Coléoptéres de lle de Cuba. Annls Soc. ent. Fr. (4) 7 : 570-616.

Cuujo, M. 1959. Coleoptera of the Loo-Choo Archipelago. Mem. Fac. Lib. Arts and Educa-
tion, Kagawa University, part 2, no. 69 : I-15.

Cuujo, M. & OunrRA, H. 1965. Elaterid beetles from Aomori Pref. Japan, collected by
Mr Kensaku Shimoyama. Mem. Fac. Lib. Arts and Education, Kagawa University, part 2,
NO 132.) 32) pp:, 2 pl:

Cogpos, A. 1964. Coléoptéres Elatérides et Throscides recoltés par M. J. Mateu dans l’Ennendi

et au Tchad. Bull. Inst. fond. Afr. noive A, 26 : 585-613, 17 figs.

1966. Nuevo género y especie de Agrypninae (Col. Elateridae). Annis Soc. ent. Fr. (N.S.)
2 : 651-653, I fig.

1970. Contribution a la faune du Congo (Brazzaville) Mision A. Villiers et A. Descarpen-
tries. Bull. Inst. fond. Afr. noive A 32 : 130-250.

CosreEs, J. P. 1782. Deliciae Cobresianae. 1. P. Cobres Biichersammlung zur Natur-
geschichte. xxxvili + 956 pp. Augsburg. The ‘Vorbericht’ of this work is dated
“Wintermonat 1781’ which suggests that the manuscript had been completed by this date.

Cotston, H.S. 1909. Peales Museum. Pop. Sci. Mon. 75 : 221-238.

Cowan, C. F. 1971. On Guérin’s Iconographie: particularly the insects. J. Soc. Biblphy
nat. Hist. 6 (1) : 18-29.

Crowson, R. A. 1961. On some new characters of classificatory importance in adults of
Elateridae (Coleoptera). Entomologist’s mon. Mag. 96 : 158-161, 7 figs.

Cuvier, G. L. C. D., Baron. 1832. Rapport de M. le Baron Cuvier sur un ouvrage de M.
Guérin intituté ‘Iconographie du régne animal’. Annis Sci. Nat. 26 : 331-334.

DEGEER, C. 1774. Mémoires pour servir a l'histoire des Insectes 4, x + 469 pp., 9 pls. Stock-
holm.

Derjean, P. F. M. A. 1833 [-35]. Catalogue des Coléoptéres de la collection de M. le Comte
Dejean. [Edit.2]. 443 pp. Paris.

Derxson, W. & SCHEIDING-GOLLNER, U. 1963-1968. Index litteraturae entomologicae
Series 2. (In progress). Deutsche Akademie der Landwirtschaftswissenschaften zu
Berlin.

DESBROCHERS DES LOGEs, J. 1875. Opuscules entomologiques (Coléoptéres). 56 pp. Gannat.

Des Gozis, M. 1886. Recherche de l’espéce typique de quelques anciens genres. 36 pp.
Montlucon.

DEYROLLE, A. 1862. In L. Maillard, Note sur l’ile de la Reunion (Bourbon), Annexe H,
Coléoptéres, 21 pp., 1 pl. Paris.

DupPERREY, L. I. 1829. Voyage autour du monde... sur... La Coquille... pendant 1822—
1825 etc. Hydrographie et Physique. Tableaux de Routes Parcourues, &C., 133 pp., I map.
Paris.

Eston, A. H. 1924. Revision of the Australian Elateridae. Part 1. Tvans. R. Soc. S.
Aust. 48 : 196-208.

1927. Revision of the Australian Elateridae. Part 2. Tvans. R. Soc. S. Austr. 51 : 350-
362.

1930. Results of Dr E. Mjéberg’s Swedish Scientific Expeditions to Australia 1910—
1913. 50. Elateridae. Avk. Zool. 22A, no. 1. 22 pp.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 285

EmpEN, F. I. van. 1945. Larvae of British beetles. 5. Elateridae. Entomologist’s mon.
Mag. 81 : 13-37.

ENGELMANN, W. 1846. Bibliotheca historico-naturalis. 786 pp. Leipzig.

Ericuson, W. F. 1834. In F. J. F. Meyen, Beitrage zur Zoologie, gesammelt auf einer

Reise um die Erde. Nova Acta Acad. Caesar. Leop. Carol. 16. Suppl. 1 : 219-284,

3 pls.

1842. Beitrage zur Fauna von Vandiemensland. Arch. Naturgesch. 8 : 83-287, 2 pls.
EscuHscHoLtz, J. F. 1829. In Thon, Entomologisches Archiv 2 (1) : 31-35. Elaterites. Jena.
Fapricius, J. F.C. 1775. Systema entomologiae. xxxii + 832 pp. Flensburgi & Lipsiae.
1792. Entomologia systematica 1 (2) 538 pp. Copenhagen.

1798. Supplementum entomologiae systematicae. 572 pp. Hafniae.

1801. Systema eleuthevatorium. 1, xxiv + 506 pp. 2,687 pp. Kiel.

FAIRMAIRE, L. 1849. Insectes de Taiti des Marquises et des iles voisines. Rev. Mag. Zool.
1 : 34-306.

1871. Notes sur les Coléoptéres recueillis par Charles Coquerel a Madagascar et sur les

cétes d’Afrique. Annls Soc. ent. Fr. (5) 1 : 29-78.

1880. Diagnoses de Coléoptéres de l’ile de la Réunion. Naturaliste 2 : 293.

1881. Essai sur les Coléoptéres des iles Viti (Fidgi). Annis Soc. ent. Fr. (6) 1 : 243-318.
1884a (April). Description de Coleoptéres Marocains. Naturaliste 2 : 446-447.

1884b (end July). Quelques Coléoptéres madécasses. Stettin. ent. Zig 45 : 273-277.
1884c. Diagnoses de Coléoptéres de 1’Afrique orientale. Annis Soc. ent. Belg. (Bull.)
28 : CXXI-CXXV.

1887. Coléoptéres des voyages de M. G. Revoil chez les Somalis et dans l’intérieur du
Zanguebar. Annis Soc. ent. Fr. (6) 7 : 69-112 (Mai), 113-186 (Aofit).

1891. Notitle. Bull. Soc. ent. Fv 1891 : LXX.

1892. Coléoptéres d’Obock. Troisiéme Partie. Revue Ent. 11 : 77-127.

1897. Matériaux pour la faune Coléoptérique de la Région Malgache (4e Note). Annis
Soc. ent. Belg. 41 : 363-406.

1901. Matériaux pour la faune Coléoptérique de la Région Malgache. (11e Note).
Revue Ent. 20 : to1—248.

1903. Matériaux pour la faune Coléoptérique de la Région Malgache. (16e Note).
Annls Soc. ent. Fr. 72 : 181-259.

FarIRMAIRE, L. & DEYROLLE, H. 1878. Description de Coléoptéres recueillis par M. L’Abbé
David dans la Chine centrale. Avmmnls Soc. ent. Fr. (5) 8 : 87-140, 2 pls.

FAIRMAIRE, L. & GERMAIN, P. (860. Coleoptera Chilensia. Revue Mag. Zool. 12 : 267-269.

FALDERMANN, F. 1835. Additamenta entomologica ad Faunam Rossicam. Fauna Persico-
Armeniaca. Nouv. Mém. Soc. imp. Nat. Mosc. 4 : 1-310, 10 pls. Reissued 1837 with a
second part loc. cit. 5 : 433 pp., 15 pls and also in two volumes under a new title, Fauna
entomologica Tvanscaucasica.

Fai, H.C. 1932. New Coleoptera 15. Can. Ent. 64 : 56-62.

Fea, L. 1896. Quattro anni fra 1 Bivmani e le tribu limitrofe. xviii + 565 pp., 195 figs,

3 maps. Milano.

FLETCHER, T. B. t919. Early stages of Agrypnus fuscipes. Bull. agric. Res. Inst. Pusa 89 :
25, 2 figs.

FLEUTIAUX, E. 1889. Contributions 4 la Faune Indo-Chinoise. ter mémoire. Cicindelidae
et Elateridae. Annis Soc. ent. Fr. (6) 9 : 137-146.

1891. Elateridae de la Nouvelle-Calédonie et dépendences. Annis Soc. ent. Fr. 61 : 387—
396.

1895a (Mars). Contributions 4 la Faune Indo-Chinoise. 15e Mémoire. Premiére
addition aux Cicindelidae et Elateridae. Amnls Soc. ent. Fr. 63 (1894) : 683-690.

1895). List de Trixagidae, Monommidae, Eucnemidae et Elateridae importés dans les
tabacs des manufactures et recueillis par M. Ant. Grouvelle. Annis Soc. ent. Belg. 39 :
157-172.

286 CoM, FS von (A AyYink

FLEUTIAUX, E. 1899. Remarques sur quelques Elatérides de Madagascar et descriptions
d’espéces nouvelles. Bull. Mus. Hist. nat. Paris 5 : 222-226.

1902 (Avril). Description de quelques Elatérides nouveaux de ma collection. Bull.
Soc. ent. Fy. 1902 : 112-116.

1902a (Mai). Descriptions de quatre espéces nouvelles du genre Agvypnus. Bull. Soc.
ent. Fy. 1902 : 162-164.

1902b (Juillet). Description de deux Adelocerva nouvelles. Bull. Soc. ent. Fr. 1902:
213-214.

1902¢ (Juillet). Note sur les Dilobitarsus Africains. Annls Soc. ent. Belg. 46 : 297-302.

1903a@ (Jan.). Contributions a la faune Indo-Chinoise. 18e Mémoire. Deuxiéme
addition aux Cicindelidae et Elateridae. Amnls Soc. ent. Fr. 71 (1902) : 569-580.

1903b (15 Nov.). Description de Coléoptéres nouveaux. Elatérides de Madagascar.
Naturaliste 25 : 263.

1903c (18 Nov.). Elateridae recueillis par le Dr Achille Tellini pendant le voyage qu’il
fit en Erythrée (octobre 1902 a février 1903). Buwill. Soc. ent. Fy. 1903 : 250-251.

1905. Voyage de M. Maurice Maindron dans |’Inde Meridionale (1901). Elateridae.
Annls Soc. ent. Fr. 74 : 319-330.

—— 1906a. Elatérides nouveaux de Madagascar. Bull. Soc. ent. Fy. 1906 : 199-200.

—— 1906b. Description de quatre Elatérides nouveaux. Bull. Soc. ent. Fy. 1906 : 211-213.
1907a (Juillet). Note synonymique. Bull. Soc. ent. Fy. 1907 : 162.

1907b (August). Révision des Elatérides du Chili. Revta chil. Hist. nat. 11 : 160-232.
—— 1908. Descriptions d’Elatérides nouveaux du Tonkin. Bull. Soc. ent. Fr. 1908 : 164.
— gir. Contribution a la faune des Hes Comores. Annls Soc. ent. Fr. 80 : 471-479.

1916. Elateridae des Philippines. Philipp. J. Sci. 11, no. 3, sec. D : 219-233.

—— 1918a (Juin). Coléoptéres Elatérides Indochinois de la collection du Muséum d’Histoire
Naturelle de Paris. Catalogue et descriptions des espéces nouvelles. Bull. Mus. Hist.
nat. Paris 24 : 205-236.

1918) (Juin). Collections recueillies par M. Maurice Rothschild dans l'Afrique Orientale.
Coléoptéres Elatérides. Descriptions des espéces nouvelles. Bull. Mus. Hist. nat.
Paris 24 : 237-241.

1918c (Aoat). Elatérides nouveaux du voyage de M. G. Babault en Afrique Orientale.
Bull. Soc. ent. Fr. 1918 : 167-170.

1918d (Décembre). Nouvelles contributions a la faune de l|’Indo-Chine Frangaise. <Annls
Soc. ent. Fr. 87 : 175-288.

1919. Insectes Coléoptéres 13. Elateridae, Trixagidae et Melasidae. In Voyage de Ch.
Alluaud et R. Jeannel en Afrique Orientale (1911-1912) Résultats Scientifiques. 119 pp.,
ipl. Paris.

1920a (Mai). Les espéces Indo-Chinois du genre Adeloceva. Bull. Soc. ent. Fr. 1920:
II2-I15. 4

1920b. (Décembre). Coléoptéres Mélasides et Elatérides des Iles Mascareignes. Mission
de M. P. Carrié (1910-1913). Descriptions des espéces nouvelles. Bull. Mus. Hist. nat.
Paris 26 : 489-499.

1921. Insectes Coléoptéres, Elateridae. In Voyage de M. G. Babault dans l Afrique
Orientale Anglaise. Résultats Scientifiques. 19 pp., 1 pl. and text illustrations, Paris. For
itinerary see Babault, 1917.

1922a (Décembre). Coléoptéres Elaterides. In Voyage de M. le Baron Maurice de Roths-
child en Ethiopie et en Afrique Orientale Anglaise (1904-1905). Résultats Scientifiques.
Animaux articulés. 2 : 288-598. Paris.
1922b. Description de Coléoptéres nouveaux (Cicindelidae, Trixagidae et Elateridae)
recueillis dans l’Angola par la Mission Rohan-Chabot. Bull. Mus. Hist. nat. Paris 28 : 44-48.
1923. Coleoptera: Melasidae et Elateridae des Seychelles et des iles voisines. Tvans R.
ent. Soc. Lond. 55 (1922) : 398-463, 1 pl.

1925a (Mai). Descriptions de deux genres nouveaux d’Elateridae. Buill. Soc. ent. Fr.
1925 : 101-103.

RECLASSIFICATION OF SUBFAMILY’ AGRYPNINAE 287

1925b (Septembre). Notes sur les genres Adelocera et Lacon. Bull. Soc. ent. Fr. 1925 :
205-207.

1926. Remarques et observations sur le catalogue des Elatérides (1er partie) de S.
Schenkling. Amnls Soc. ent. Fr. 95 : 91-112.

1927. Les Elatérides de l’Indochine Frangaise (catalogue raisonné). Faune Colon. Fr.
1 : 53-122, 2 pls. Paris.

1931. Contributions a la faune d 1|’Indochine frangaise. Bull. Soc. ent. Fr. 36 : 73-76.

1932a (Mars). Elateridae nouveaux de Madagascar. Bull. Soc. ent. Fr. 37 : 49-51.
1932b (June). Contribution a l’étude de la faune du Mozambique. Voyage de M. P.
Lesne (1928-1929). 6e Note. Coléoptéres, Elateridae. Mems Estud. Mus. zool. Univ.
Coimbra Ser. 1, no. 55, 23 pp.

1932c (July). Contributions 4 la faune Malgache. (Coléoptéres, Elatérides). Bull. Annis
Soc. v. ent. Belg. 72 : 152-157.

1932d (Novembre). Description de trois Coléoptéres nouveaux. Bull. Soc. ent. Fr.
37 : 228-230, 2 figs.
1932e (15 Décembre). Jkemarques sur quelques espéces de Motschulsky. Miscnea ent.
34: 79-80. [Note. As the result of a printer’s error the pages are numbered 71 and 72.
Correction, loc. cit. : 80.]

1932f (21 Décembre). Elatérides nouveaux de Madagascar. Bull. Soc. zool. Fr. 57:
450-456.

1933. Entomological investigations on the Spike disease of Sandal (10). Melasidae &
Elateridae. Indian Forest Rec. 18, part 7. 16 pp.

1934@ (February). Schwedisch-Chinesische wissenschaftliche Expedition nach dem
nordwestlichen Provinzen Chinas. Insekten gesammelt von Dr D. Hummel 1927-30.
39. Coleoptera 7. Elateridae. Avk. zool. 27A, no. 19 : 15-21.

1934) (March). Contributions a la faune Malgache. Coléoptéres, Elatérides. Bull. Soc.
zool. Fr. 59 : 49-68.

1934¢ (July). Elateridae nouveaux des Iles Philippines. Proc. Hawaii ent. Soc. 8:
(1933) 477-486.

: 1934d (Aoait). Descriptions d’Flatérides nouveaux. Bull. Soc. ent. Fr. 39 : 178-185.

1934e€ (Novembre). Eucnemidae et Elateridae nouveaux. Bull. Annis Soc. ent. Belg.

74 : 363-370.

| 1935a@ (Mars). Reévision des genres Agvaeus Candéze et Tvachylacon Motschulsky. Annls
Soc. ent. Fr. 104 : 11-39.

— 1935b (April). Coleoptera 5. Elateridae. In Mission scientifique de l’Omo. 2. Zoologie.

Fascicle Ir : 193-216, 10 figs, 1 pl. Paris.

1935c (June). Description d’Eucnemidae et Elateridae nouveaux provenant du Coryndon
Memorial Museum de Nairobi. Ji E. Africa Uganda nat. Hist. Soc. 12 : 90-117.
1940a (Mars). Elatérides nouveaux. Bull. Annls Soc. ent. Belg. 80 : 89-104.
1940b (Mai). Un nouvel Elatéride de Bornéo. Bull. Soc. ent. Fr. 45 : 32.
1941a (Mars). Les Elatérides de l’Indo-Chine frangaise. Huitiéme et derniére partie.
Annls Soc. ent. Fy. 109 (1940) : 19-40.
1941b (Mars). Elatérides nouveaux de Indochine frangaise. Bull. Soc. zool. Fr. 65
(1940) : 184-189.

—— 194Ic (1Avril). Tableau des genres et sous-genres des Octocryptitae et des Agrypnitae.
Revue fr. Ent. 8 : 42-48.

1941d (25 Avril). Mission P. Lepesme, R. Paulian et A. Villiers au Cameroun.
Coléoptéres, Elatérides. Bull. Mus. natn. Hist. nat. Paris (2) 13 : 38-44.

1942. Entomological results from the Swedish expedition to Burma and British India:
Elateridae collected by R. Malaise. Avk. Zool. 33A, no. 18, 24 pp.

1944. Les Octocryptus Candéze. Revue fr. Ent. 10 : 145-148, 4 figs.

1945. Historique de la collection d’Elatérides de Candéze, actuellement au Musée Royal
d’Histoire Naturelle de Bruxelles. Revue fr. Ent. 12 : 79-81.

1947. Revision des Elatérides del’Indo-Chine francaise. Notes Ent. chin. 11 : 233-420.

288 C. M. F. von HAYEK

FuEss.i, J.C. 1778. Etwas tiber Voet’s Kaferwerk. Magazine Liebh. Ent. 1 : 1-7o.

Gauan, C. J. 1900. On a collection of insects and arachnids made in 1895 and 1897 by
Mr C. V. A. Peel, F.Z.S. in Somaliland, with descriptions of new species. Coleoptera.
Proc. zool. Soc. Lond. 1900 : 21-33, 1 pl.

GEMMINGER, M. & HaRoLpD, E. von. 1869. Catalogus Coleopterorum, 5 : 1347-1604.
Monachii.

GERMAR, E. F. 1824. Insectorum spectorum species novae aut minus cognitae, descriptionibus
illustvatae. 1. Coleoptera. xxiv + 624 pp., 2 pls. Halae.

1840. Bemerkungen tiber Elateriden. Z. Ent. (Germar) 2 : 241-279.

1844. Bemerkungen tiber Elateriden. Z. Ent. (Geymar) 5 : 133-102.

1848. Beitrage zur Insectenfauna von Adelaide. Linn. ent. 3 : 153-247.

GERSTAECKER, G. E. A. 1856. Necrolog von Klug. Sétettin. ent. Zig 17 : 225-237.

1871. Beitrage zur Insektenfauna v. Zanzibar. III. Avch. Naturgesch. 37 : 42-86.

1873. Baron C. C. von der Decken’s Reisen in Ost Afrika in 1859-61. 3, Abth. 2. xvii +
542 pp., 18 pls. Leipzig & Heidelberg.

GIRARD, C. 1970. Note sur quelques Coléoptéres Elatérides de Madagascar. Mem. Orstom,
no. 37 : 25-29.

GMELIN, J. F. 1790. C.a.Linné... Systema Naturae...Editio decima tertia ... Cura J. F.
Gmelin, 1, pt. 4 pp., 1517-2224. Lipsiae.

GoLBacH, R. 1953. Revision de las Adelocerinae de la Argentina. Acta zool. lilloana 13:

205-306, 2 pls, 5 figs.

1968. Pyrganus nov. gen. Acta zool. lilloana 22 (1967) : 197-199.

— 1969a. Anotaciones sobre el género Meristhus Cand. Act zool. lilloana 25 : 137-142.
— 1969b. Anotaciones sobre el género Lacon Cast., 1836. Acta zool. lilloana 25 : 151-160.
—1970a. Sinonimia de los géneros Acrocryptus Cand. y Hewxaulacus Cand. Acta zool.
lilloana 25 : 305-307.

1970b. Alotipo de Lacon (Cornilacon) faiymairet (Cand.). Acta zool. lilloana 25 : 327-329.
GRIFFIN, F. J. 1932. On the dates of publication and contents of the parts of Westwood,

J. O. Introduction to the modern classification of insects, 1838-1840. Proc. R. ent. Soc.
Lond. 6 : 83-84.

1936. On the dates of publication of Motschulsky (V. de), ‘Etudes entomologiques’
I-IX, 1853-1862. Ann. Mag. nat. Hist. (10) 17 : 256-257.

GUERIN-MENEVILLE, F. E. 1829. JIconographie du végne animal de G. Cuvier, 2, Insectes,
pl. 12. For dating of this plate see p. 59, and Cowan, 1971.

1830. In Duperry, ed. Voyage autour du monde sur ...la Coquille pendant les années
1822-1825. Atlas, Insectes, pl. 2.

1838. Crustacés, arachnides et insectes. Jn Duperry, ed. Voyage autour du monde
sur... La Coquille pendant les années 1822-1825. Zoologie 2, p. 2, div. 1. [111] iv—xii +
pp. 9-319. Paris.

1844. Iconographie du régne animal de G. Cuvier. 3 [made up of fascicles 46—50 of text].
xvi + 567 pp. Paris.

GYLLENHAL, L. 1808. Insecta svecica ... Classis 1. Coleoptera sive Eleuthevata. 1. vii +
572 pp. Scaris.

1817. Appendix ad C. J. Schénhery synonymiam insectorum 1, part 3, 266 pp. Scaris.
Hacen, H. A. 1857. Nachricht tiber das Kaferwerk von Joh. Euseb. Voet. Stettin. ent. Zig
18 : 405-409.

1862. Bibliotheca entomologica 1, 566 pp. Leipzig.

HELLER, K. M. 1937. Coleoptera Javana, mit Beriicksichtigung der ihnen nahestehenden
Formen anderer Herkunft. Stettin. ent. Zig 98 : 51-78, 6 figs, 1 pl.

Hersst, J. F. W. 1779. Beschreibung und Abbildungen einiger theils neuer, teils noch

nicht abgebildeter Insecten. Beschdft. berlin. Ges. naturf. Fr. 4 : 314-326, 1 pl.

1784. Verzeichnis seiner Insectensammlung. Fortsetzung. Avch. Insectengesch. 5:

73-151, 6 pls.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 289

1806. Natursystem alley bekannten in- und auslandischen Insecten. Kafer. 10. viii +
285 pp., 19 pls. Berlin.

HEYDEN, L. von & Kraatz, G. 1882. Kafer um Samarkand, gesammelt von Haberhauer.
Dt. ent. Z. 26 : 297-338.

Hore, F. W. 1831. Synopsis of the new species of Nepaul insects in the the collection of
Major General Hardwicke. Jn J. E. Gray, Zoological Miscellany, part 1 : 21-32. London.

1838. Observations on succinic insects. Part 2. Tvans. ent. Soc. Lond. 2 (1837-1840) :

46-58, 1 pl. For date of publication see Wheeler, 1912.

1843a. Description of the Coleopterous insects sent to England by Dr Cantor from
Chusan and Canton, with observations on the entomology of China. Ann. Mag. nat. Hist.
(1) 11 : 62-66. This is a shortened version of Hope, 1845. See also Wheeler, 1912.
1843b. Onsome new insects from western Africa. Ann. Mag. nat. Hist. (1) 11 : 364-369.

— 1845. Onthe entomology of China, with descriptions of the new species sent to England
by Dr Cantor from Chusan and Canton. Trans. ent. Soc. Lond. 4: 4-17, 1 pl. A short
version of this paper appeared in 1843.

Horton, A. 1949. Kdferkunde fiir Naturfreunde. 292 pp., 21 pls. Frankfurt am Main.

Horn, G. H. 1871. Descriptions of new species of Elateridae of the United States. Tvans.
Am. ent. Soc. 3 : 299-324, 1 pl.

1875. Synonomical notes and description of new species of North American Coleoptera.

Trans. Am. ent. Soc. 5 : 126-156.

1894. The Coleoptera of Baja California. Proc. Calif. Acad. Sci. (2) 4 : 302-449, 2 pls.
Horn, W. & Kaute, I. 1935-1937. Uber entomologische Sammlungen, Entomologen &
Entomo-Museologie. Ent. Beih. Berl.-Dahlem 2-4. vi + 536 pp., 38 pls, 3 text-fig.
Horn, W. & SCHENKLING, S. 1928-1929. Index litteraturae entomologicae. 4 vol. 1426 pp.

Berlin.

HORSFIELD, T. & Moore, F. 1857. A catalogue of the lepidopterous insects in the museum of
The Hon. East-India Company 1. 278 + iv + 11 [+7] pp., 6 pls. London.

Huser, F. & J. 1940. Studien uber die Biologie der Elateriden. Mitt. miinch. ent. Ges.
30 : 343-396.

Hystop, J. A. 1917. The phylogeny of Elateridae based on larval characters. Ann. ent.
Soc. Am. 10 : 241-263.

1921. Genotypes of the Elaterid beetles of the World. Pyoc. U.S. natn. Mus. 58 : 621-

680.

ILLINGwortTH, J. F. 1921. Natural enemies of sugar-cane beetles in Queensland. Bull.
Bur. Sug. Exp. Stns Qd Div. Ent. 13. 47 pp., 8 pls.

JACQUELIN DU VAL, C. 1859. Genera des Coléoptéres d'Europe. 3, 1859-63. 456 pp.,
100 pls. Paris. Livraisons 78-87 (pp. 1-136) appeared in wrappers dated 1859.

JAGEMANN, E. 1944. De Elateridarum speciebus novis. Sborn. ent. Odd. nav. Mus. Praze
21-22 : 332-335.

James, E. 1823. Account of an expedition from Pittsburgh to the Rocky Mountains performed
in the years I819 and 1820 under the command of Major S. H. Long. 3 vols, with map and
illustrations. London.

Junk Catalogue see Schenkling, S.

Kayser, C. G. 1841. Index locupletissimus libvorum . . . Vollstandiges Biicher-Lexicon
enthaltend alle von 1750 . . . in Deutschland und in den angrenzenden Ldandern dedvuckten
Biicher .. . bearbeitet... von C. G. Kayser, etc. Thlg 7, 566 pp. Leipzig.

KIESENWETTER, H. von. 1863. Jn Erichson, Naturgeschichte der Insecten Deutschlands.
Erste Abteilung. Coleoptera 4. 745 pp. Berlin.

Kinnear, N. B. 1948. Obituary, J. R. D. Bell. J. Bombay nat. Hist. Soc. 48 : 167-171.

Kirk, H. B. 1922. Biological notes on Elateridae and Melasidae. Ent. News 33 : 236-240.

Kirkatpy, G. W. 1902. Publication date of the Rhynchotal part of the ‘Voyage of the
Coquille’. Entomologist 35 : 316.

Kirscu, T. F. W. 1865. Beitrage zur Kaferfauna von Bogota. Berl. ent. Z. 9 : 40-140.

aly

290 C. M. F. von HAYEK

Kisuu, T. 1957. Elateridae of Nachi, Japan, with description of a new species. Akitu

6 : 84-89.

1958. Snappers from Kyftishu District, Japan, collected by Prof. Takashi Shirozfi. Ent.

Rev. Japan 9 : 27-32.

1959. Elateridae of Amami-Oshima collected by Prof. T. Shirézu. Akitu 8 : 57-66.
1961. Elateridae of Is. Tsushima. ‘The Snappers of Island (2)’. Bull. Heian High

School Kyoto 5:56 pp., 11 pls. [Reprint, original not seen. ]

1964. Elateridae of Islands Awa-shima, Hegura-jima and Nanatsu-jima. ‘The Snappers
of Islands (4)’. Bull. Heian High School, Kyoto 8 : 38 pp., 2 pls, I map.

Kiue, F. 1833. Bericht tiber eine auf Madagascar veranstaltet Sammlung von Insecten der
Ordnung Coleoptera. Phys. Math. Abh. K. Akad. Wiss. Berlin, Separate preprint issued
1833, 135 pp., 5 pls [see Klug, 1834 loc. cit.: 215 and Kayser, 1841 : 520]. 1834, loc.cit.:
gI-223. The preprint dated 1833 appears to have been generally available. It was used
by Candéze, 1857 and is quoted by Schenkling, 1925. Ihavenotseenacopy. The pagina-
tion appears to correspond thus: 1833 : 3 = 1834 : 91 to 1833 : 133 = 1834: 220. 1834:
121-123 contain the captions to the plates. In the main body of the text of the present
work the page number of the 1834 issue is given in parentheses after the 1833 reference to
assist workers not having access to the 1833 preprint.

1855. Diagnosen neuer Coleopteren aus Mossambique. Ber. Akad. Wiss. Berlin 1855:

643-660.

KNUuLL, J. N. 1949. New Coleoptera with notes. Ohio J. Sci. 49 : 102-104.

KoeEnic, E. 1887. Neue Elateriden und Bemerkungen tiber bekannte Arten. Tvudy russk.
ent. Obshch. 21 : 353-354.

Ko.sBe, H. J. 1889. Bericht iiber die von Herrn Dr R. Bittner in Gebiete des unteren
Quango und Kongo gesammelten Coleopteren. Stettin. ent. Ztg 50 : 113-133.

Koirar, V. 1844. Coleoptera. In C. F. von Huegel, Kaschmir und das Reich dev Siek 4
part 2 : 497-564. Stuttgart.

KONTKANEN, P. 1964. On the heterogeneity of the genus Hypnoidus Steph. sensu auct.
(Coleoptera, Elateridae). Suomen hyént. Aikak. 30 : 45-49.

KueEster, H.C. 1845. Die Kafer Euvopas.2 [ii + 200pp.],2pls. Nutrnberg.

1850. Die Kafer Europas. 20 : [ii + 200 pp.], 2 pls. Niirnberg.

LACORDAIRE, T. 1857 (June). Histoive naturelle des insectes. Geneva des Coléoptéves. 4.
579 pp. Paris. This volume is first recorded in the Bibliographie de la France for 27
June, 1857, (2) 1 no. 26 : 302, no. 5737.

Lane, M.C. Some generic corrections in the Elateridae. 4. Proc. ent. Soc. Wash. 55 : 86-89.

Laporte, F. L. DE, see Castelnau.

LaTREILLE, P. A. 1809. Insectes de 1 Amérique équinoxiale recueillis pendant le voyage de
MM. de Humboldt et Bonpland. In Humboldt, F. H. A. von & Bonpland, A. J. A.
Voyage aux régions equinoxiales du nouveau continent, fait en 1799-1804 etc. (2) 2, livraison
4, 197-283, pls 15-18. The date 1811 recorded by the Schenkling catalogue refers to the
re-issue of this work.

1827. In Cailliaud, F. Voyage a Méroé, au Fleuve Blanc etc. 4, Description des

Insectes: 271-292. Atlas2:pl.2. Paris.

1829. Jn Cuvier, G.L.C. F. D., Le végne animal. Nouvelle édition. 4, xxvii + 584 pp.

Paris.

1834. Distribution méthodique et naturelle des genres de diverses tribus d’insectes

Coléoptéres de la famille des Serricornes. Annis Soc. ent. Fr. 3 : 113-170.

Lea, A. M. 1920. Descriptions of new species of Australian Coleoptera. Proc. Linn. Soc.
N.S.W. 45 : 375-403.

LeConTeE, J. L. 1853. Revision of the Elateridae of the United States. Tvans. Am. phil.
Soc. 10 : 405-508.

1857. Report upon the insects collected on the survey. Reports of the explorations and
surveys for a railroad from the Mississippi River to the Pacific Ocean. 9, No. 1, 72 pp., 2 pls.
(editor) 1859a. The complete writings of Thomas Say on the entomology of North America.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 291

With a memoir of the author, by George Ord. 2 vol., New York. Not examined, but

believed to be identical with the 1869 edition except for the title page.

1859b. Additions to the coleopterous fauna of northern California and Oregon. Proc.

Acad. nat. Sci. Philad. 11 : 281-292.

1866. Additions to the coleopterous fauna of the United States. No. 1. Proc. Acad.

nat. Sci. Philad. 18 : 361-394.

(editor) 1869. American entomology. A description of the insects of North America
by Thomas Say. 2 vols. xxiv + 412 pp. and v + 814 pp., 13 pls. New York.

LEsNE, P. 1940. Remarques sur divers Elatérides. Bull. Soc. ent. Fr. 45 : 33-37.

Lewis, G. 1894. On the Elateridae of Japan. Ann. Mag. nat. Hist. (6) 13 : 26-48.

1896. On new species of Coleoptera from Japan, and notices of others. Ann. Mag.
nat. Hist. (6) 17 : 329-343.

LINNAEUS, C. VON, 1758. Systema naturae. toth ed. 1, [ii] 824 pp. Holmiae.

1761. Fauna suecica, sistens animalia ... etc. Ed. alt. (xlvi) + 578 pp. 2 pls. Stock-

holmiae.

1766. Systema naturae. 12th ed. 1, pp. 1-532. Holmiae.

1767. Ibidem, 2, pp. 533-1327 (+36). Holmiae.

MacLeay, W. J. 1872. Notes on a collection of insects from Gayndah. Tvans. ent. Soc.
N.S.W. 2 : 239-318. Note: Chadwick (1964, J. ent. Soc. Australia (N.S.W.) 1: 4) states

that this volume was published in 1873. However, the Zoological Record for 1873 records

MacLeay’s paper. Derksen & Scheiding-Géllner (1968 : 22) also record the date as 1872.

1888. The insects of the vicinity of King’s Sound, N.W. Australia. Pyvoc. Linn. Soc.

N.S.W. (2) 3 : 1227-1246.

MarsEvL, S. A. DE, 1870-71. Descriptions de Coléoptéres nouveaux. Abeille 6 : 368-389.

1872-73. Répertoire des Coléoptéres d’Europe décrits isolément depuis 1863. (2° partie).
Abeille 9 : 1-448.

Matsumura, S. 1911. Beschreibungen von am Zuckerrohr Formosas schadlichen oder
nutzlichen Insecten. Mem. Soc. y. ent. Belg. 18 : 129-150.

MELSHEIMER, F. V. 1806. A catalogue of insects of Pennsylvania. vi-+ 60 pp. Hanover,
York County.

MENETRIES, E. 1832. Catalogue vaisonné des objects de zoologie vecueillis dans un voyageau
Caucase et jusqu aux frontiéves actuelles de la Perse. 271 + xxxlipp. St Pétersbourg.
MENKE, A. S. 1962. The dates of publication of Palisot de Beauvois’ ‘Insectes Recueilles

[sic] en Afrique et en Amérique.’ Ann. Mag. nat. Hist. (13) 5 : 701-702.

Micnot, E.C. 1969. See Arnett, Mignot & Smith, 1969.

Miwa, Y. 1927. Descriptions of new species of Japanese Elateridae. Insecta matsum. 2,

No. I 2 12=22, r pl.

1929. Elateridae of Formosa. Tvans. nat. Hist. Soc. Formosa 19 : 225-246.

1931. Elateridae of Formosa. Tvans. nat. Hist. Soc. Formosa 21 : 72-98.

1933. Elateridae in the collection of the Entomological Laboratory, Kyushu Imperial

University. Mushi 6 : 25-31.

1934. The fauna of Elateridae in the Japanese Empire. Rep. Dep. Agric. Govt res.

Inst. Formosa No. 65, 289 pp., 9 pls.

Mocuuv’sku, V. See Motschulsky.

MoTscHULSKY, V. DE. 1854. Diagnoses de Coléoptéres nouveaux, trouvés par MM. Tatarinoff
et Gaschkewitsch aux environs de Pékin. Etudes entomologiques 2 : 44-51. Helsingfors.

1858. Insectes des Indes Orientales. Etudes entomologiques 7 : 20-122. Helsingfors.

1861a. Insectes du Japon. I. Coléoptéres. Etudes entomologiques 9 (1860) : 4-27.

Helsingfors. For date of publication see Griffin, 1936.

1861b. Essai d’un catalogue des insectes de I’Ile Ceylan. Bull. Soc. Nat. Moscou
34 : 95-155.

Moucuet, J. 1949. Deux Elasmosomus nouveaux de madagascar. Mém. Ins. scient.
Madagascar (A) 3 : 113-115, I fig.

Murray, A. 1868. List of Coleoptera received from Old Calabar on the west coast of

292 C. M. F. von HAYEK

Africa. Ann. Mag. nat. Hist. (4) 2: 91-111. Murray published a volume with the same
title in 1878. Though the title page states that the work is ‘From the Annals and Magazine
of Natural History, 1857-71’ the text does not always correspond exactly with that of the
separate parts. Murray, 1868 : 91-111 = 1878: 137 (Metriorhynchus)-158 (end of
Chrysodema).

MusGrRaveE, A. 1932. Bibliography of Australian entomology 1775-1930. Royal Zoological
Society of N.S.W. Sydney. 380 pp.

NAKANE, T. 1964. Iconographia insectorum Japonicorum colove natuvali edita. 2nd ed. 2,
433 pp., 192 pls. Hokuryu-kan, Tokio.

NAKANE, T. & Kisui1, T. 1555. Entomological results from the scientific survey of the

Tokara Islands. 1. Coleoptera, Elateridae. Bull. Osaka Mus. nat. Hist. No. 2, 8 pp.,

2 pls.

1956. On the subfamilies of Elateridae in Japan. Kontyu% 24 : 210-206.

Negsoiss, A. 1956. A check list of Australian Elateridae. Mem. natn. Mus. Vict. No. 22

part 2. Ent. series 1, 75 pp.

1961. Additions and corrections to the check list of Australian Elateridae. Mem.

natn. Mus. Vict. No. 22, part Io, Ent. series 3, 29 pp.

NisHio, Y. 1960. On the female copulatory organs of some beetles. Kontyu% 28 : 142-145,
6 figs.

OutrRA, H. 1954a (February). Notes on some generic names of Japanese Elateridae. New

Ent. Ueda 3. Nos. 2-3, 10 pp. [In Japanese].

1954b (December). Notes on some species of Elateridae from Shikoku. Gensei 3, Nos

I-2 : 14, I fig. [Note: this periodical, which is not listed in the World List of Scientific

Periodicals, 4th edition, is published by the Entomological Laboratory, Agricultural

Faculty, Kochi University. }

1957. Studies on the morphology of the Elaterid larvae of Japan (3). New Ent. Ueda

6, No. 4 : 4-8.
1958. Studies on the morphology of Elaterid larvae of Japan (2). Kontyi 26 : 83-87,
2 pis.

1959. Some larval Elateridae of Kochi Prefecture. Gensez No. 9 : 65-66, figs. [See
Ohira, 1954).]

1962. Morphological and taxonomic study on the larvae of Elateridae in Japan. 179 pp.,
61 pls. Japan. Published by the author.

1966a (March). Notes on some Elaterid-beetles from the Ryukyu Islands. Bull. Jap.
ent. Acad. 2,no.2 : 7-12. [Note: not recorded in the World List of Scientific Periodicals,
4th edition. Published by the Biological Laboratory, Nanzan University, Showa-ku,
Nagoya. ]

1966b (September). Notes on some Elaterid-beetles from Formosa (1). Kontyi# 34 : 215-
D222.

1967a (April). Notes on some Elaterid-beetles from Formosa (4). Kontyi 35 : 55-59.
1967b (May). The Elateridae of the Ryukyu Archipelago (1). Tvans. Shikokt ent. Soc.
9 : 95-106.

1968a (April). Some Elaterid-beetles collected by the late Mr K. Ohbayashi in Japan.
Ent. Rev. Japan 20 : 73-77.

1968b (November). Notes onsome Elateridae-beetles from Formosa V. Kontyt 36 : 363—
368.

1969a (February). The Elateridae of the Ryukyu Archipelago (4). Bull. Aichi Gakuget
Univ. 18 (Nat. Sci.) : 89-103.

1969b (March). Notes on some species of Elateridae of the Izu Islands. Tvans. Shikoku
ent. Soc. 10 : 41-48.

1969c (April). The Elateridae of the Ryukyu Archipelago (7). Bull. Jap. ent. Acad.
4 : 27-32.

1970a. Anew Elaterid-beetle from the Bonins. Gemnsei 21: 9-11. (See Ohira, 19540.)

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 293

1970b. A list of the Elaterid-beetles from South Asia in the Hungarian Natural History

Museum. Parts 1-V. Anymls hist.-nat. Mus. natn. hung. 62 : 207-243.

1971a. Notes on some Elaterid-beetles from south-east Asia. Bull. Aichi Gakugei Univ.

22 (Nat. Sci.) : 225-236.

1971b. New or little-known Elateridae from Japan, XIII. Kontyt% 39 : 40-41.
1972c. On some Elaterid-beetles from Japan (VI). Kontyu% 39 : 63-66.

Otivier, G. A. 1790. Entomologie ou histoive naturelle des insectes, Coléoptérves 2, No. 31,
54 pp.,8pls. Paris.

Orp, G. 1859. See Leconte, J. 1859a.

ORELLANA, B. 1937. Revision de los Elateridos chilenos. El genero Adelocera Latreille.
Revta chil. Hist. nat. 40 (1936) : 57-60.

PaisoT DE BEeauvois, A. M. F. J. 1805. Insectes vecueillis en Afrique et en Amérique, dans
les royaumes d’Oware et de Benin, a Saint-Domingue et dans les Etats Unis pendant les années
1786-1797. xvi-+ 276 pp., 90 pls. Paris. For date of publication see Menke, 1962.

Panzer, G. W. F. 1793. Johan Euseb Voet’s Beschreibungen und Abbildungen hartschaaligter
Insekten, Coleoptera, Linn. Aus dem Original getreu tibersetzt, mit der in selbigem fehlenden
Synonymie etc. Thl. 2. 134 pp., 15 pls. nos. 23-48. Erlangen.

1795. Deutschlands Insectenfaune, oder entomologisches Taschenbuch fiiv das Jahr 1795.

[vill + 24] 370[2] pp., 13 pls. Nurnberg.

1796. Faunae insectorum germanicae initia odey Deutschlands Insecten. Heft 34, No. 12.

Niirnberg.

1801. Ibidem. Heft 76, No.1, No. 4. Niirnberg.

Pascok, F. P. 1885. List of the Curculionidae of the Malay Peninsula. Annali Mus. civ.
Stor. nat. Giacomo Doria (2) 22 : 201-331.

PAvuLiAN, R. 1947. Un nouvel Elateride de Cote d’Ivoire. Bull. Soc. ent. Fr. 52 : 7, 1 fig.

PaYKULL, G. 1800. Fauna suecica; Insecta. (Coleoptera) 3, 459 pp. Uppsala.

PérincuEy, L. 1892. Third contribution to the South African coleopterous fauna. Tvans.
S. Afr. phil. Soc. 6, part 2 : 1-94.

Perty, J.A.M. 1830. InSpix, Delectus animalium articulatorum quae in itineve per Brasiliam
annis 1818-1820... collegerunt J]. B. de Spix...et...C.F.P. Martius. Digessit. descripsit,
pingenda curavit M. Perty etc. Fasc. 1, 111 + 44 + 60 pp., 12 pls. For dates of publica-
tion see Isis, Jena 1832 : 137.

Puiviprl, F. H. E. 1861. Species quaedam novum Coleopterorum Chilensium. An. Univ.

Chile 1861 : 742-744. Not examined, not available in BMNH library.

1887. Catalogo de los Coleopteros de Chile. An. Univ. Chile 71 : separate section of

190 pp.

Puitipri, R. A. 1873. Chilenische Insekten. Stettin. ent. Zig 34 : 296-316; 2 pls.

Puitipri, A. (sic, in error for F.}.H. E.& R.A. 1860. Coleoptera nonnulla nova Chilensia
praesertim Valdiviana. Stettin. ent. Ztg 21 : 245-251. Dohrn (loc. cit. : 252) states that
the species without the reference to “Ph’. were described by F. H. E. Philippi.

Pic, M. 1928. Mélanges exotico-entomologiques. 51, 36 pp. Moulins.

Pore, R. D. 1960. A revision of the species of Schizonycha Dejean (Col: Melolonthidae)
from southern Africa. Bull. By. Mus. nat. Hist. (Ent) 9 : 63-218, 14 pls.

QUEDENFELDT, [F. O.] G. 1886. Verzeichniss der von Herrn Major a D. von Mechow in
Angola und am Quango-Strom 1877-1881 gesammelten Buprestiden und Elateriden.
Berl. ent. Z. 30 : 1-38, 1 pl.

QUELLE, F. 1932. Elateriden-Studien 2. Coleopt. Zbl. 5 : 202-216.

1955. Elateriden-Studien 4. Dt. ent. Z. (N.F.) 2 : 225-229.

Racusa, E. 1881. Coleotteri nuovi o poco conoscuiti della Sicilia. Naturalista sicil. 1 : 5-9,

ie pl.

1gi1. Coleotteri nuovi o poco conoscuiti della Sicilia. Riv. coleott. ital. 9 : 190-108.

Razoumowsky, G. DE. 1789. AHistoive naturelle du Jovat et de ses envivons, etc. 1, xvi + 32 pp.,

3 pls. Lausanne.

294 Cc. Ms BF. von HAYEK

ReicuHE, L. & SauLcy, F.DE. 1856 (Nov.). Espéces nouvelles ou peu connues de Coléoptéres

recueillies par M. F. de Saulcy dans son Voyage en Orient. Annls soc. ent. Fr. (3) 4 : 353-384.

1857 (Jan.). Continuation of the above paper. Amnmnls Soc. ent. Fr. (3) 4 : 385-422, pl. 12.

REICHENSPERGER, A. 1915. Myrmekophilen und Termitophilen aus Natal und Zululand
gesammelt von Dr I. Tragardh. Meddelanden fran Géteborgs Musei Zoologiska Aldelning
5, 20 pp., in Goteborgs K. Vetensk.-o. vitterhSamh. Handi. 16.

REITTER, E. 1905. Bestimmungs-Tabelle der europdischen Coleopteren, 56 Heft. Elateridae,
Elaterini, Subtribus Athouina aus der palaearctischen Fauna. Verh. naturf. Ver. Brunn
43, Abhandlungen: 3-122.

1911. Fauna germanica. Die Kafer dey Deutschen Reiches. 3 : 463 pp., 48 pls.

Ritsema, C. 1881. Description of a new species of the Coleopterous family Elateridae.

Notes Leyden Mus. 3 : 29-30.

1891. Alphabetische Naamlijst der beschreven soorten van het Melolonthiden-genus
Apogoma Kirby. Tijdschr. Ent. 34 : xcii—xcvii.

Rovusat, J. 1928. Adelocera fasciata L. A. cejkai N. A. Cas. ésl. Spol. ent. 25 : 46.

SAHLBERG, J. 1903. Coleoptera mediterranea et rosso-asiatica nova et minus cognita, maxima
exparte itineribus annis 1895-1896 et 1898-1899 collecta. Ofvers. finska VetenskSoc.
Férh. 45, no. 10, 40 pp.

SAKURAI, K. 1942. On the Wireworms injurious to agricultural and horticultural plants in
Hokkaido. Kontyu% 16, no. 1 : 1-11, I fig.

SALVADORI, T. 1878. Descrizioni di una nuova specie di Uccello del genere Chalcopsittacus,
Bp. e note intorno ad altre specie di Uccelli della Nuova Guinea. Atti Accad. Sci. Torino
13 : 309-316.

Say, I. 1825. Descriptions of new American species of the genera Buprestis, Tvachys and
Elater. Ann. Lyceum nat. Hist. 1, part 2: 249-268.

1839. Descriptions of new North American insects, and observations on some already
described. Tvans. Am. phil. Soc. (N.S.) 6: 155-190. This paper, which was published
posthumously (Say died in October, 1834), is a continuation of one published in the same
journal five years before (1834 (N.S.) 4 : 409-470). There is no indication that the whole
or part of either paper has been published before but LeConte (1869 : 521, footnote)
states that part of Say’s work, as far as Paganus (1834 : 431), was published in the Dis-
seminator [of useful knowledge containing hints to the youth of the U.S. from the School of
Industry, New Harmony, Indiana] from June to August 1830 and another part from the
beginning to E. obesus (1839 : 168) in an 8vo pamphlet with a title page dated New Har-
mony, Indiana, 1829-33. This is probably the pamphlet of 65 pp. published in New
Harmony, 1829-33 recorded by Englemann (1846 : 551). Horn and Schenkling (1928 :
1050) state that the whole work was first published as a pamphlet of 81 pp. dated 1829-33
by the Disseminator of Useful Knowledge, New Harmony. Carus and Englemann
(1861 : 519) record a work of the same name and length published in New Harmony,
1830-1834.

I have been unable to obtain a copy of the pamphlet{s] and as far as can be ascertained
there are no copies of the Disseminator in this country. In view of the uncertainty of the
date of publication of the three species (Elater obtectus, awrovatus and avita) considered in
this work, the generally accepted date of 1839 has been retained for the present.
Scuaum, H. R. 1853. Necrolog von Ernst Friedrich Germar. Stettin. ent. Ztg 14 : 375-390.
SCHENKLING, S. 1925. Coleopterorum catalogus, auspiciis et auxilio W. Junk. Pars 8o,

Elateridae 1,263 pp. Berlin.
1927. Ibidem, Pars 88, Elateridae 2, pp. 265-636. Berlin.
SCHIQDTE, J. G. 1865. Danmarks Buprestes og Elateres. Naturh. Tidsskr. (3) 3 : 441-456,

I pl.
ScumipT, F. J. 1832. Leptodivus Hochenwarthi und Elater Grafii beschrieben. Faunus
1 : 83-86.
SCHOENHERR, C. J. 1817. Synonymia insectorum 1, pt. 3. xi + 506 pp., 2 pls. Stockholm
and Skara.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 295

SCHRANK, F. von P. 1781. Enumeratio insectorum Austriae indigenorum. 548 pp., 4 pls.
Augustae Vindelicorum.

ScHwWARZ, O. 1891. Zwei neue Elateriden von Syrien und Madagascar. Dt. ent. Z. 1891 :

151-152.

1893. Adelocera kvaatzi Schwarz nov. spec. Dt. ent. Z. 1893 : 192.

1894. Neue Elateriden. Dt. ent. Z. 1894 : 145-150.

1898a. Beschreibung neuer Elateriden. Dt. ent. Z. 1898 : 129-156.

1898b. Elateriden aus Usambara (Ost-Afrika). Dt. ent. Z. 1898 : 183-192.

1898c. Aenderung der Gattungsnamen Enoplodeves und Craspedonotus. Dt. ent. Z.

1898 : 414.

1899. Neue von Herrn v. Bennigsen in Deutsch Ost-Africa gesammelte Elateriden.

Dt. ent. Z. 1899 : 74-80.

1900. Neue palaarctische Elateriden. Dt. ent. Z. 1900 : 98-112.
1901. Verzeichniss der von Drn Horn auf Ceylon gesammelten Elateriden nebst

Beschreibung neuer Arten. Dt. ent. Z. 1901 : 17-32.

1902a (January). Neue Elateriden. Df. ent. Z. 1901 : 311-336.

19026 (ist July). Neue Elateriden. Stettin. ent. Zig. 63 : 194-316.

1go2c (July). Neue Elateriden aus Australien. Dt. ent. Z. 1902 : 113.

1g02zd. Neue Elateriden aus Siid Amerika, besondersaus Peru. Dt. ent. Z.1902 : 129-152

1g0ze. Aenderung der Speciesnamen lineatus Schw. (Lacon) und variegatus Schw.

(Hypnoidus). Dt. ent. Z. 1902 : 155.

1902f (October). Neue Elateriden aus dem tropischen Asien, den malayischen Inseln

und den Inseln der Siidsee. Dt. ent. Z. 1902 : 305—350.

1903a. Neue Elateriden aus Afrika und Madagascar. Dt. ent. Z. 1903 : 357-376.
1903b. Neue Elateriden aus Neu-Guinea und Australien. Dt. ent. Z. 1903 : 377-3098.
1903¢c. Die von Herrn Prof. Yngve Sjéstedt in Kamerun gesammelten Elateriden,

Eucnemiden und Throsciden. Ark. Zool. 1 : 43-53.

1904. Diagnosis of a new species of Elateridae in the Colombo Museum. Spolia zeylan.

2: 46.

1905a. Neue Elateriden aus der Malayischen Zone. Dt. ent. Z. 1905 : 257-266.

1905b. Neue Elateriden aus Madagaskar und Afrika. Dt. ent. Z. 1905 : 273-2092.

1906-1907. In Wytsman, Geneva Insect. Fasc. 46A-C, Coleoptera, Fam. Elateridae.

370 pp., 6 pls.

1908a (August). Neue Elateriden aus Afrika und Madagaskar. Societas ent. 23 : 73.

1908) (1 September). Ibidem, loc. cit. : 82-83.

1g08c (15 September). Ibidem, loc. cit. : 93.

1908d (1 Oktober). Ibidem, loc. cit. : ror.

1g08e. In Sjostedt, Wissenchaftliche Ergebnisse dey Schwedischen Zoologischen Expedition
nach dem Kilimandjaro, dem Meru und den Umgebenden Massai-steppen Deutsch-OstA frikas,
1905-1906. 1, part 7, no. 6, Elateridae., pp. 57-66, The whole work was not published
until 1910 but the Elaterid section is recorded in the Zoological Record for 1908.

Scopoti, G. A. 1763. Entomologia carniolica, exhibens insecta Carnioliae indigena etc.
[xxxvi] + 420 pp., 43 pls. Vindobonae.

ScuDDER, S. H. 1897. Dates of issue of some of Boisduval’s works. Psyche, Camb.
8 : 153-154.

SEIDLITZ, G. 1872. Fauna baltica. Die Kafer (Coleoptera) der Ostseeprovinzen Russlands.
1872-1875. xlii + 560 pp. Dorpat. This work appeared in several parts. For dates of
publication see preface.

1888. Fauna baltica. 2nd ed. 1887-1891. [10] + lvi+ 818 pp., 1 pl. K6nigsberg.
This work appeared in six parts. For dates of publication see preface.

SHARP, D. 1877. On the Elateridae of New Zealand. Ann. Mag. nat. Hist. (4) 19 : 396-413
and 469-487. Note: this work was also issued as a separate with pagination I-37.

SHarP, D. anp Scott, H. 1908. Fauna hawaiiensis, 3, part 5 : 367-579, 4 pls. Cambridge.

HITT |

Poidd

FETT

296 Co Me EH) von (EVA 2k

SHERBORN, C. D. & GRIFFIN, F. J. 1934. On the dates of publication of the natural
history portions of Alcide d’Orbigny’s ‘Voyage Amerique méridionale’. Ann. Mag. nat.
Hist. (10) 13 : 130-134.

SHERBORN, C. D. & Woopwarp, B. B. igor. Notes on the dates of publication of the

natural history portions of some French voyages. Ann. Mag. nat. Hist. (7) 7 : 388-392.

1906. Notes on the dates of publication of the natural history portions of some French
voyages. ‘Voyage autour du Monde sur la Coquille pendant 1822-25 par L. J. Duperry’,
etc. A Correction. Ann. Mag. nat. Hist. (7) 17 : 335-336.

SmiTH, E.H. 1969. See Arnett, Mignot & Smith, 1969.

SoLieR, A. J. J. 1851. In Gay, Historia fisica y politica de Chile. Zoologia, 5, 563 pp.

Paris & Santiago.

1854. Ibidem, Aflas. 2 Coleoptera. 32 pls.

Sotsky,S.M. 1871. Coléoptéres de la Sibérie orientale. Hovae Soc. ent. voss. 7 (1870) :334—

406. Note: from 1865 (see Horae etc, 1866, 3 : xx) ? the contents of the Horvae differ from

those of the Tvudy russk. ent. Obshch. Cf World List of Scientific Periodicals, 4th ed.

1881. Coléoptéres nouveaux ou peu connus de l’Empire Russe et des pays limitrophes.

Trudy russk. ent. Obshch. 12 (1800-1881) : 231-265. [In Russian and Latin.}

SZOMBATHY, C. 1909. Notes sur quelques Elaterides exotiques. <Avnmnls hist.-nat. Mus. natn.
Hung. 7; 118-124, 4 figs.

Tuomson, C.G. 1859. Skandinaviens Coleoptera 1. [2] + 290 pp. Lund.

1864. Skandinaviens Coleoptera 6. 385 pp. Lund.

1868. Skandinaviens Coleoptera 10. 358 pp. Lund.

TSCHEREPANOV, A. I. 1957. ([Click-beetles of westeyn Siberia (Coleoptera, Elateridae), &c.}.
382 pp. Novosibirsk. [In Russian, with German summary.] [Academie fur wissenschaften

USSR],
VaN Dyke, E. C. 1943. New species of west American Coleoptera. Pan. Pacif. Ent.
19 : 41-52.

Van ZWALUWENBURG, R. H. 1945. Notes on the Samoan Elaterid Beetles with descriptions

of two new species. Occ. Pap. Bernice P. Bishop Mus. 18 : 95-101.

1947. Some Elaterid beetles from Australia and New Guinea. Proc. Hawai. ent. Soc.

13 : 109-115.

1957. Coleoptera. Elateridae. Insects Micronesia 16, No. 2 : 56 pp., 12 figs.
1959. Some type designations with notes on Pacific Elateridae. Pacif. Insects 1,

No. 4 : 347-414.

1966. <Adelocera nesiotes, new name for Adelocera cognata 1957, preocc. Pacif. Insects

8, No. 1 : 289.

Vauri£, P. A. 1960. A revision of the genus Diplotaxis (Coleoptera, Scarabaeidae,
Melolonthinae). Part 2. Bull. Am. Mus. nat. Hist. 120, article 2 : 161-434, 229 figs.
Vinson, J. 1963. Sur l’apparition nouvelle 4 Maurice d’un scarabée nuisible, Oryctes rhino-

cevos Linné. Revue agric. sucyr. Ile Maurice 42, No. 3 : 173-180. Addendum.

VoET, J. E. 1781. Catalogus systematicus Coleopterorum. 1 (1769-1781) 88 pp., 48 pls. La
Haye. Page 85 (sheet L3) appears to have been published between 1778 and 1781 (Hagen,
1857 : 406) but the exact date is unknown. Horn & Schenkling (1929 : 1266) state that
at the time of Voet’s death in 1778, sheets A-L (=pp. 1-88) had already appeared. Horn
& Schenkling refer to Cobres (see below) but this author does not record the dates of
publication of the separate sheets. Fuessli (1778 : 4, probably published early in the
year, see note on the back cover of part 1) records only 9 sheets, pp. I-72 [=sheets A-—I].
By 1781 (Cobres, 1782 : 230, see p. 285) 88 pages [=sheets A—L] and 48 plates but no title
page had appeared.

WALKER, F. 1866. Coleoptera. In Lord, The naturalist in Vancouver Island and British
Columbia. 2 : 309-334. London.

WEBER, F. 1801. Observationes entomologicae, continentes novorum quae condidit generum
characteres, et nuper detectarum specierum descriptiones. xii + 116 pp. Kiliae.

RECLASSIFICATION OF SUBFAMILY AGRYPNINAE 297

WeEstwoop, J. O. 1838. An intvoduction to the modern classification of insects. 2, Synopsis.
Coleoptera, 1-44. For date of publication see Griffin, 1932.

WHEELER, G. 1912. On the dates of the publications of the Entomological Society of

London. Trans. R. ent. Soc. Lond. (1911) : 750-767.

ZIMSEN, E. 1964. The type material of I. C. Fabricius. 656 pp. 2 pls. Copenhagen.

ZINCKEN, J. L. T. F. (also known as Sommer), 1818. Knochs Tod. Magazin Ent. (Germar)
3 : 458-460.

In addition to the Times Index-Gazetteer of the World, 1965, London (see p. 10) the following
gazetteers have been consulted in the course of this work.

Lippincott’s Gazetteer of the World. Philadelphia, 1886.

Lippincott’s Gazetteer of the World. Philadelphia & London, 1922.

Longmans’ Gazetteer of the World. London, 1895.

Stieler’s Hand-Atlas, Namenverzeichnis. Gotha, 1925.

U.S. Navy Department, Hydrographic Office. Gazetteers.

U.S. Board of Geographic Names. Gazetteers prepared in the Office of the Geography Depart-
ment of the Interior, Washington.

INDEX

This index includes the names of all the genera and species included in the
Agrypninae as defined in the Notes on the Key on p. 5. Species not known to me
are recorded with the reference to the original description and the original generic
attribution. Only the original spelling is shown for adjectival specific names whose
Invalid names are

endings are liable to change according to the generic gender.

in italics.
the main entry is given in bold type.

aberdarensis Fleutiaux, 119

abbreviatus Candéze, 94, 95

aberrans Candéze, 53, 68

abreptus Candéze, 119

abstrusus nom. n., 118, 120

acerbus Candéze, 120

acervatus Candéze, 120

Acrocryptus Candéze, 8 (key), 92, 268

aculeatus Candéze, 120

acuminatus Fleutiaux, 241

acuminipennis Fairmaire, 120

acutuangulus Fleutiaux, 121

adanensis Jagemann, 12, 21

adelaidae Blackburn, 121

Adelocera Latreille, 6 (figs), 7(key), 13, 54,
268

Adelocera Latreille sensu auct., 15, 52, 54, 86,
87, 114, 116, 268

Adelocerina Blackwelder, 12
U

Where a name appears more than once in the text the reference to

Adelocerinae Fleutiaux, 12

Adelocerini Buysson, 12

adeloceroides Candéze, 121

adspersa Candéze, 1874, 76

adspersa Fleutiaux, 57

adspersulus Klug, 1855 : 647, Dicrepidius
adspersus Candéze, 1857, 22

adspersus Herbst, 114, 240

adustus Elston, 118, 121

aequalis Candéze, 1857, 241, 242, 248, 254
aequalis Candéze, 1900, 122

aethiopicus Candéze, 23, 185

afflictus Candéze, 122

Aganolacon Ohira, 13, 16, 268

Agraeus Candéze, 13, 15, 16, 268
Agrypnidae Fleutiaux, 12

Agrypnides Candéze, 12

Agrypnina Thomson, 12

Agrypninae Fleutiaux, 12

298 Cc. M. F. von HAYEK

Agrypnini Kiesenwetter, 12

Agrypnitae Fleutiaux, 12

Agrypnites Jacquelin du Val, 12

Agrypnus Eschscholtz, 6(figs), 7(figs, key),
15, 113, 118, 231, 235, 240, 268

Agrypnus Eschscholtz sensu auct., 113, 240,
269

akidiformis Candéze, 113, 122

Alaotypus Schwarz, 52, 54, 268

Alaus, 240

alberti nom. n., 122

albicans Chevrolat, 81

albisparsus Candéze, 123

albitactus Candéze, 123

alboguttatus MacLeay, 1888 : 1238, Lacon

albomaculatus Miwa, 123

albopictus Candéze, 123

alboscutatus Candéze, 124

alboseminatus Kolbe, 156

Aliteus Candéze, 240

alluaudi Candéze, 102

alluaudi Fleutiaux, 1919, 23

alluaudi Fleutiaux, 19346, Centrostethus, 124

alluaudi Fleutiaux, 1934d, Prolacon, 13, 15,
16, 32

altaica Candéze, 56

alternans MacLeay, 1872 : 251, Lacon, 227

alternatus schwarz, 124

amamiensis Miwa, 1934 : 248, Paralacon

Amaurus Burmeister, 240

Amaurus Castelnau, 240, 269

ambiguus Schwarz, 19050 : 279, Lobotarsus

amplicollis Boheman, 124, 280

Anacantha Solier, 93, 94, 268

Anaspasis Candéze, 4, 5, IOI

anathesinus Candéze, 12

Anathesis, 12

andersoni Blackburn, 125

andrewsi Candéze, 243

andrewsi Fleutiaux, 23

angularis Schwarz, 115, 116, 125

angulicollis Candéze, 125

angulicollis Fairemaire, 232

angulosus Candéze, 1857 : 117, Lacon

angusta Fleutiaux, 125

angustata Philippi, 1861 : 743, Adelocera, 100

angustata Sahlberg, 89

Anius Candéze, 8, 85, 268

Anomala, 272

antennata Lewis, 56

antennatus Candéze, 250, 257

anthracina Candéze, tgoo : 1, Adelocera

antiguus Candéze, 125, 130

apicalis Candéze, 236

apicalis Schwarz, 1898, 109

apicalis Schwarz, 1905, 102

apodixus Candéze, 126

applanatus Elston, 126

aquilus Elston, 1927 : 360, Lacon

arabicus Candéze, 243

arbitrarius Elston, 126

Ayrchontas Goezis, 113, 268

Archontoides Cobos, 113, 115, 117, 269

arctior Candéze, 126

arenicola Nakane & Kishi, 127

argentatus Candéze, 1874, 56, 91

argentatus Candéze, 1893, 127

argillaceus Solsky, 127, 140, 147

aristatus Champion, 127

arizonae Candéze, 243

armatus Candéze, 128

Aynettia Golbach, 53, 55, 268

asper Candéze, 128

aspersus Schwarz, 54, 57, 114

asperulatus Candéze, 128

assus Candéze, 128

ater Philippi, 92, 93

atricolor MacLeay, 1888 : 1236, Lacon

atomarius Fabricius, 52, 53, 54, 56, 76, 77,
II4, 240

attenuatus Candéze, 243

attermima Candéze, 57

attonitus Candéze, 244

Aulacon Arnett, 53, 55, 268

auricollis Schwarz, 63

auroratus Say, 57, 80, 276

aurosquamosa Jagemann, 1944 : 323, Adelo-
cera

aurulenta Candéze, 58, 70, 295

australasiae Dup. nomen nudum, 261

australis Candéze, 244

avitus Say, 58, 295

babaulti Fleutiaux, 1918, 241, 244
babaulti Fleutiaux, 1944, 266
baccatus Fleutiaux, 24

bacillus Candéze, 102

badeni Candéze, 1878, 129
badeni Candéze, 1889, 245
badius Elston, 1927 : 350, Lacon
baibaranus nom. n., 129

bakeri Fleutiaux, 1916, 129
bakeri Fleutiaux, 1935, 24
bartoni Fleutiaux, 245

basalis Fleutiaux, 129
beauchenii Fleutiaux, 24
beccarii Candéze, 129

INDEX 299

beninensis Candéze, I1

benitensis nom. n., 130

bellator Elston, 1927 : 354, Lacon

bergeali Girard, 130

bhasini Fleutiaux, 24

bicolor Fleutiaux, 58

bicornis Candéze, 95

bicostata Schwarz, 76

bidens Fabricius, 93, 95

bidentata Fleutiaux, 130

bidivisus Candéze, 130

bifasciatus Schwarz, 1902b : 312, Lacon

biforatus Candéze, 1885) : 55, Lacon

bifoveatus Candéze 245

bigener Elston, 131

bigranosus Schwarz, 1905) : 276, Lacon

biguitatus Candéze, 236

bimaculatus Schwarz, 1902c : 113, Lacon

bimarginatus Schwarz, 131

binodulus Motschulsky, 118, 131, 209

binus Candéze, 1889 : 73, Lacon

bipapulatus Candéze, 132, 193

bipunctatus Candéze, 132, 243, 245

bipunctatus Heller, 1937 : 54, subspecies of
scobinula Candéze

bipunctatus Schwarz, 132

birmanicus Candéze, 188

birmanicus Fleutiaux, 25

bituberculatus Fleutiaux, 103

bituberculus Schwarz, 103

blackburni Fleutiaux, 132

blairi Fleutiaux, 133

bocandei Candéze, 260

bohemanni Dejean nomen nudum, 232

boildieus Candéze, 146

boninensis Ohira, 1970a : 11, Alaotypus

borneensis Fleutiaux, 25

brachychaetus Kollar, 133

Brachylacon Motschulsky, 13, 15, 16, 268

bradshawi Candéze, 246

brasiliensis Castelnau, 80

breve Golbach, 133

brevicornis Fleutiaux, 57

brevicornis LeConte, 87, 88

brevipennis Elston, 1927 : 360, Lacon

brevipennis Schwarz, 1903c : 44, Lobotarsus

brevis Candéze, 133

brightensis Blackburn, 133

brulerievi Desborcheres des Loges, 63

brunneipennis Candéze, 134

brunneiventris Schwarz, 103

brunneus Lewis, 25

Bruyantius Fleutiaux, 51, 268

bullatus Carter, 1939 : 301, Lacon

bulwensis Fleutiaux, 58
buyssoni Jagemann, 1944 : 332, Lacon

caffer Candéze, 1857, 108, 109

caffer Candéze, 1881, 134

calabarica Candéze, 58

calamitosus Candéze, 134

caliginosa Boisduval, 18, 19, 20, 21, 134, 135,
173

caliginosa Guérin-Méneville, 134, 136

caliginosus Candéze, 252

cambodiensis Fleutiaux, 25

Candanius nom. n. 8(key), 85, 268

candezei Desbrochers des Loges, 59

candezei Fleutiaux, 1895, 137

candezei Fleutiaux, 1919, 264

candezei Fleutiaux, 1935, 46

candida Fall, 59

canescens Candéze, 137

capensis Fleutiaux, 52

carbonarius Schrank, 76

cardiophoroides Fleutiaux, 110

cardoni Candéze, 266, 267

cariei Fleutiaux, 137

carinensis Candéze, 59

carinicollis Schwarz, 138

carinulatus Candéze, 121, 138

cariosus Candéze, 96

castaneipennis Candéze, 138

castaneus Elaston, 1927 : 358, Lacon

castelnaui Candéze, 1882 : 11, Lacon

castelnaui Candéze, 1889, 59

catulus Candéze, 3, 26

catulus Candéze, 9, 26

cavicollis LeConte, 88

Cavicoxum Pic, 13, 15, 269

cejaki Roubal, 1928: 46, aberration of
fasciata Linnaeus

Centrosthethus Schwarz, 113, 115, 116, 268

cervinus Erichson, 116, 138

chabannei Guérin-Meéneville, 53, 59

Chalcolepidius, 78

chappuisi Fleutiaux, 110

chapuisti Candéze, 74

charcus Boheman, 43

chilensis Solier, 60

chinensis Hope, 272

chrysoprasa Herbst, 1806 : 18, Elater

churakagi Ohira, 19710 : 40, Alaotypus

cincta Candéze, 60

cinctipes Germar, I13, 115, 118, 139

cinerascens Candéze, 139

cineraceus Elston, 139

300 C.-M. F. von HAYEK

cinereus Candéze, 246

cinnamomeus Candéze, 113, 116,118, 139,147

cithareus Candéze, 1893) : 9, Lacon
coarctatus Candéze, 140

Coccinella, 273

coctus Candéze, 140, 172

coeca Candéze, 60

coenosus Candéze, 149

coenosus Hope, 114, 140, 149
cognata Candéze, 61

cognatus Van Zwaluwenburg, 141
Colaulon Arnett, 113, 116, 117, 268
collisa Candéze, 1889 : 70, Adelocera
collisus Candéze, 1891, 141
colombianus Candéze, 96

colonicus Candéze, 141, 219

colorata Fleutiaux, 142

communis MacLeay, 1888 : 1238, Lacon
commutabilis Elston, 142

compactus Candéze, 1882 : 10, Lacon
Compsolacon Reitter, 113, 116, 118, 268
comptus Candéze, 142, 279

confusus Candéze, 170

confusus Fleutiaux, 1935, Agrypnus, 241, 246

confusus Fleutiaux, 1935, Trachylacon, 26
consobrinus Candéze, 142

consors Candéze, 142

conspersa Gyllenhal, 14, 87, 88, 114
conspersus Dejean, nomen nudum, 256
conspiciendus Elston, 1924, 143
conspiciendus Elston, 1927, 151
conspurcatus Candéze, 143

constellata Fleutiaux, 61

constrictus Ritsema, 16, 26
controversa Hope, 272

coomani Fleutiaux, 1927, Agraeus, 29
coomant Fleutiaux, 1927, Lacon, 72
coomani Fleutiaux, 1944, 118, 267
cordicollis Candéze, 143

cordipennis Candéze, 143

coreanus Kishii, 131

Cormilacon Golbach, 53, 55, 268
cornutus Candéze, 103

corrosus Candéze, 96

Corymbites, 54, 264

corvinus Candéze, 1882 : 12, Lacon
costicollis Candéze, 121, 144
costipennis Germar, 144

cotesi Candéze, 1889 : 73, Lacon
crassiusculus Candéze, 246
crassiventris Schwarz, 247

crassus Candéze, 1874 : 88, Lacon
crenatus Germar, 20, 145

crenatus Klug, 20, 54, 144, 190, 196, 274

crenatus Schwarz, 190

crenicollis Ménétries, 20, 54, 113, 145, 221
cribrata Candéze, 61

cribratus Blanchard, 91

cribrosus Eschscholtz, 72, 114, 136
cristata Fleutiaux, 61

cristatus Horn, 233, 235

crocopus Hope, 272

cruentus Olivier, 1790 : 45, Elater
cruentatus Elston, 1927 : 251, Lacon
crux Philippi, 86, 94, 95, 97, Ioo
Cryptohypnus, 18

Cryptolacon Nakane & Kishi, 113, 116, 269
Cryptotarsus Philippi, 92, 93, 268
cuneatus Candéze, 62, 95

cuniculus Candéze, 37

curtus LeConte, 146, 204

cuspidatus Klug, 113, 115, 116, 146, 226
cylindricus Candéze, 115, 223
cylindricus Fleutiaux, 103
cylindriformis Fleutiaux, 27
cylindripennis Fleutiaux, 147

cypri Baudi, nomen nudum, 12

Danosoma Thomson, 7(fig, key), 87, 240, 268

davidi Fairmaire, 147

dealbatus Candéze, 147

deboulayi Candéze, 147

decipiens Boheman, 27, 280

decoratus Candéze, 148, 177

decovatus Schwarz, 113, 177

defectus Candéze, 148

delagrangei Buysson, 62

delamarei nom. n., 102, 104

delesserti Candéze, 148

Delox Quelle, 87, 268

demissus Candéze, 27

denticollis Fleutiaux, 148

denticornis Fleutiaux, 83

depressus Candéze, 1874, 130, 149

depressus Candéze, 1882 : 14, Tilotarsus

depressus Schwarz, 130

desjardinsii Candéze, 149

desquamatus Candéze, 149

dewalquu Candéze, 247

deyrollei Candéze, 97

deyrollei nom. n., 149

difficilis Lewis, 28

dilatatus Fleutiaux, 28

dilaticollis Fleutiaux, 150

Dilobitarsus Latreille 8(key), 86, 93, 94, Lot,
268

diodesmoides Motschulsky, 236

INDEX 301

Diphyaulon Arnett, 53, 55, 268
discedens Candéze, 1878, 150
discedens Candéze, 1892, 15, 29
discoidea Weber, 62

distincta Fleutiaux, 62
distinctus Fleutiaux, 33
divaricatus Candéze, 115, 150
divergens Fairemaire, 247
doherti Fleutiaux, 29

dorcinus Candéze, 150

dorsalis Candéze, 62

dorsalis Schwarz, 110

drusa Marseul, 63

dubius Candéze, 63, 94, 117
duboulayi Candéze, 147
duchoni Schwarz, 1905a : 257, Adelocera
duodevigintispilota Hope, 273
duplex Candéze, 132

duplex Blackburn, 151
duplicatus nom. n., 110
duporti Fleutiaux, 35

edmundi Quelle, 104

Eidolus Candéze, 8(key), 90, 268

Elasmosomus Schwarz 8 key, 94, 101, 109,
268

elegans Fleutiaux, 111

elliensis Candéze, 151

ellipticus Candéze, 151

eloini Candéze, 97

elongatus Carter, 1939 : 302, ?>Myrmodes

elongatus Sharp, 5

elstoni Neboiss, 151

Enoploderes Falderman, 113

Enoploderes Schwarz, 113, 116, 268

ereptus Candéze, 247, 248

eveptus Fleutiaux, 248

eveptus Janson, nomen nudum, 247

erinaceus Candéze, 1874, 236

erinaceus Candéze, 1897 : 12, Meristhus, 231,
232

eucalypti Blackburn, 118, 152

excavatus Fleutiaux, 29, 30

exiguus Candéze, 30

eximus Candéze, 152, 278

expansa Fleutiaux, 63

extvactus Fleutiaux, 265

fairmairei Candéze, 1882 : 3, Anacantha, 94,
95

fairmairei Candéze, 1889, 152

fallaciosus Fairmaire, 248

fallax Fairmaire, 152

falsarius Candéze, 153

falsus Fleutiaux, 30

farinensis Blackburn, 153

farinosus Candéze, 153

farinosus Elston, 1927 : 335, Lacon
fasciatus Candéze, 92

fasciatus Fabricius, 53

fasciatus Linnaeus, 14, 53, 89, 114, 240
fasciculatus Schwarz, 1902, ror, 104
fasciculatus Schwarz, 1905, III
fasciolata Candéze, 5

fasciolatus MacLeay, 1888 : 1239, Lacon
fastigiatus Schwarz, 153

fatuus Candéze, 154

feai Fleutiaux, 31

feneriva Fleutiaux, 154

feralis Candéze, 154, 168

fergusoni Elston, 154

feroculus Candéze, 32

feroculus Fleutiaux, 26

ferrantei Buysson, III

ferrugineus Candéze, 154
ferruginipes Fleutiaux, 155

fex Candéze, 155

fibrinus Candéze, 155, 205

fictus Candéze, 155

filiformis Candéze, 104

filum Candéze, 105

flammula Blanchard, 59, 60
flavescens Candéze, 155

flavipes Candéze, 175

fleutiauxi nom. n., Adelocera, 32
fleutiauxi nom. n., Lanelater, 248
fleutiauxi, Paulian, 94, 95, 102, 104
fleutiauxi Schwarz, 1902, 64
fleutiauxi Schwarz, 1906, 102, 105
floridanus Knull, 1949 : 102, Lacon
foedus Candéze, 118, 156
formosanus Bates, 156

fovmosanus Miwa, 113, 116, 129
foveata Candéze, 64

foveicollis, Cobos, 156

foveicollis MacLeay, 1888 : 1237, Lacon
frenchi nom. n., 156

froggatti MacLeay, 1888 : 1234, Lacon
fryi Fleutiaux, 32

fuliginosus Candéze, 113, 157
fulvastra Fleutiaux, 157

fulvicollis Motschulsky, 13, 15, 33, 50
fulvipennis Fleutiaux, 64
fulvisparsus Candéze, 157

funebris Solsky, 64

funestus Candéze, 248

302 C. M. F. von

furunculosus Candéze, 130, 157

fuscipes Fabricius, 114, 241, 249

fuscipes Olivier, 257

fuscus Erichson, 1841 :
158

fuscus Fabricius, 14, 157, 203

fuscus Melsheimer (nomen nudum), 65

fuscus Quedenfeldt, 204

fusiformis Candéze, 249

106, Cratonychus,

gabonensis nom. n., 158
ganglbauri Schwarz, 65
gayndahensis MacLeay, 1872
gazagnairei Candéze, 158
geminatus Candéze, 158
geographica Candéze, 52, 54, 65
gestroi Candéze, 249

gibberosus Candéze, 159

gibberosus Fleutiaux, 34

gibbosus Schwarz, 19030 : 379
gibbus Candéze, 159

giluus Candéze, 253

glirinus Candéze, 159

goudotii Candéze, 159

gracilentus Schwarz, 160

gracilis Blackburn, 113, 115, 118, 221
gracilis Candéze, 1874, Dilobitarsus, 97
gracilis Candéze, 1874, Lacon, 160
gracillimus Candéze, 5, 85, 86

graeca Candéze, 64, 65

grafil Schmidt, 1832 : 83, Elater
gragatensis Szombathy, 160

grandis Hope, 249

granulatus MacLeay, 161

granulosus Eschscholtz, 114, 124
gratiosa Fleutiaux, 90

grisea Guérin-Méneville, 134, 136
grisea, Schwarz, 1900 : 98, Adelocera
griseopilosa Fleutiaux, 161

griscens Candéze, I61

guttatus Candéze, 161, 180
gutturosus Fairmaire, 250

gypsatus Candéze, 162

: 250, Lacon

hackeri Elston, 1927 : 357, Lacon

haedulus Candéze, 162

hageni Candéze, 250

hamai Ohira, 1967 : 99, subspecies of
scutellaris Candéze

hamatus Candéze, 18936 : 8, Lacon

hamatus Schwarz, 1903a : 360, Lobotarsus

hastatus Candéze, 162

HAYEK

haterumana Ohira, 163

Hemicleus Candéze, 8(key), 101, 108, 268

herzi Koenig, 163

hexagonus Candéze, 163

Hexaulacus Candéze, 92, 93, 268

hilaris Candéze, 34

himalayensis Jagemann, 1944 : 334, Comp-
solacon

himerensis Ragusa, 256

hispidulus Candéze, 118, 164

holosericeus Candéze, 164

Homeolacon Blackburn, 113, 115, 268

horni Schwarz, 164

hottentota Candéze, 164

hova Fleutiaux, 165

humeralis Schwarz, 105

humilis Erichson, 165

hydropictus Dup. nomen nudum, 176

hydropicus Fairmaire, 1880 : 293, Lacon

hypnicola Kishi, 165

Hypnoidus, 18

illimis Horn, 166

imerensis Binaghi, 256

impressicollis Elston, 166

impressicollis Say, 65

impressicollis Schwarz,
bitarsus, 95

impressus Candéze, 166

inaequalis Candéze, 66

inaequalis DeGeer, 1774 : 148, Elater

inaequalis Fleutiaux, 166

incerta Fleutiaux, 167

incisus Fleutiaux, 1927 : 59,
mucronatus Candéze, 255

incisus Fleutiaux, 1940, 34

incommodus Fleutiaux, 66

incompta Kraatz, 66

incultus MacLeay, 1888 : 1235, Lacon

incurvata Fleutiaux, 167

indecorus Candéze, 1892c, Meristhus, 232

indicus Fleutiaux, 34

indosinensis Fleutiaux, 167

inductus Candéze, 167

indutissimus Candéze, 167

inflata Candéze, 1857, 67

inflatus Candéze, 1897, 168

infuscatus Klug, 244, 250, 257, 264

inopinus Candéze, 98

inops Candéze, 168

inordinatus Fleutiaux, III

insignitus Candéze, 1874, Lacon, 168

insignitus Candéze, 1874, Meristhus, 234

1902d : 129, Dilo-

variety of

INDEX 303

insularis Candéze, 1874, 169

insularis Candéze, 1889 : 70, Adelocera
insularis Fairmaire, 242

insulsus Candéze, 169

intermedius Schwarz, 170
interpunctatus Klug, 170

irroratus Candéze, 98

irroratus Klug, 20, 54, 170, 171
ishigakiensis Ohira, 47

jacquieri Candéze, 67

jansoni Fairmaire, 171

javana Candéze, 1893 : 6, Adelocera
javanus Candéze, 251

jeanneli Fleutiaux, 1919, 109, 111
jeanneli Fleutiaux, 1935, 110
jeanneli Mouchet, 105

jeanvoinei Fleutiaux, 67

judaicus Reiche & Saulcy, 251
judex Candéze, 163, 172
jurulosus Candéze, 172, 223

kawamurae Miwa, 1929 : 230, Lacon, 141
kenyensis Fleutiaux, 172

kikuchii Miwa, 1929 : 226, Adelocera
kilimandjarensis Fleutiaux, 35
kinangopa Fleutiaux, 173

Kobulacon Chujo & Ohira, 53, 55, 268
kokeilii Kiister, 1845 : no. 6, Lacon
koshunensis Miwa, 219

kosunensis Ohira, 219

kraatzi Schwarz, 59

labeculatus Candéze, 251

labiosus Candéze, 173

lacerta Candéze, 61

lacertosus Candéze, 251

lacrymosus Candéze, 173

Lacon Castelnau, 7(fig.), 8(key), 52, 86, 87,
268

Lacon Castelnau sensu auct, 13, 18, 20, 53,
54, 113, 116, 231, 268

laconoides Candéze, 12

laconoides Fleutiaux, 98

laconoides Schwarz, 67

lakhoni nom. n., 173

lameyi Fleutiaux, 174

Lanelater Arnett, 6(fig.), 7(fig, key), 240, 269

laosensis Ohira, 252

laoticus Fleutiaux, 68

lapideus Candéze, 174

lateralis Schwarz, 174

laticollis Candéze, 1857, Adelocera, 53, 68,
78, 94

laticollis Candéze, 1857, Lacon, 118, 174

laticollis Hope, 252

Latilacon Golbach, 53, 55, 268

latior MacLeay, 252

latiusculus Candéze, 175

latus Candéze, 175

lavaudeni Fleutiaux, 175

laxatus Candéze, 176

laxus Candéze, 35

Lepidelatey Smith, 53, 55, 268

lepidoptera Panzer, 14, 68

lepidopterus Chevrolat, 232

lepidotus Palisot de Beauvois, 231, 232, 235

Lepidotus Asso, 52

Lepidotus Stephens, 52, 53, 55, 268

leprosus Candéze, 176

lepturus Say, 66

leucaspis Candéze, 176

lezeleucii Candéze, 127, 176

libellus Candéze, 1893c : 170, Lacon

lignarius Candéze, 98

limbatus Candéze, 1857, Ocneus, 52, 54, 69

limbatus Candéze, 1857, Opatelus, 91, 92

Jimosus Candéze, 177

lindensis Blackburn, 177, 270

linearis Candéze, 1857, 90

linearis Candéze, 1865, 69

linearis Fleutiaux, 105

linearis Schwarz, 105

lineatellus MacLeay, 1888 : 1236, Lacon

lineatus Candéze, 32, 164

lineatus Schwarz, 164

lithophila Candéze, 69

litiginosus Candéze, 1874 : 73, Lacon

lobicollis Motschulsky, 30, 35

Lobitarsus Fleutiaux, 113, 116, 268

Lobotarsus Schwarz, 113, 115, 116, 268

lolodorfensis nom. n., 177

longicollis Candéze, 1889, 252

longicollis Candéze, 1897 : 11, Meristhus, 232

longicollis Schwarz, 112

longicornis Champion, 53, 69

longicornis Fleutiaux, 177

longicornis Gahan, 253

longipennis Schwarz, 178

longus Fleutiaux, 178

lopezei Fleutiaux, 1934, Adelocera, 178

lopezei Fleutiaux, 1934, Agrypnus, 241, 253

lucasseni Candéze, 37

luctuosus Candéze, 1893c : 171, Lacon

luctus Fleutiaux, 178

Ludius, 272

304 C2. M. BY von TAY EK

lupinosus Candéze, 178, 190
luridus Fabricius, 114, 253
lustratus Candéze, 179
luteipes, Hope, 272

lutosus Candéze, 118, 179
luzonica Candéze, 70
luzonicus Fleutiaux, 37

macarthuri Fleutiaux, 37

macgregori Fleutiaux, 179

macleayi Candéze, 1882 : 11, Lacon

macroderus Candéze, 179

maculata LeConte, 70

maculatus Brangsik, 180

maculatus MacLeay, 162, 180, 216

maculatus Schwarz, 216

maculicollis Gerstaecker, 254

maculipennis Schwarz, 1903a@ : 361, Lobo-
tarsus

maculosus Candéze, 1874, 180, 227

maculosus Candéze, 1893, 37

maculosus MacLeay, 1888 : 1238, Lacon, 227

madecassa Fleutiaux, 180

madida Candéze, 54, 70

madurensis Candéze, 1893c : 171, Lacon

maenamii Ohira, 19696 : 41, Alaotypus

maullavdi Deyrolle, 175

maindroni Fleutiaux, 267

major Candéze, 180

majusculus Candéze, 1897 : 8, Hemicleus

maklini Candéze, 70

mamillatus Candéze, 1857, 180

mamillatus Candéze, 1865, 71

mansuetus Blackburn, 180

mannerheimii Candéze, 13, 15, 37

marginatus Candéze, 181

marginipennis Schwarz, 1903@ : 380, Lacon

marmorata Candéze, 94, 95, 98

marmorata Fabricius, 14, 56, 71, 98

marmoratus Candéze, 1874, 118, 152, 181,
184, 230

marmovatus Candéze, 1878, 230

massiet Fleutiaux, 212

masterst var C. Candéze, 247, 248

mastersi MacLeay, 241, 252, 255

mausoni nom.n., 54, 71

Mecynocanthus Hope, 113, 115, 268

mekrani Candéze, 1889 : 8, Adelocera

melancholicus Candéze, 1890, I81

melancholicus Candéze, 1892, 223

Melanotus, 14, 158

Melonontha, 272

meridionalis Fleutiaux, 182

Meristhus Candéze, 7 (key), 231, 235, 269

mexicana Candéze, 71

microcephalus Motschulsky, 13, 15, 16, 38

microcephalus Motschulsky sensu Kishii, 28

minicephalus Ohira, 1970) : 236, Brachyla-
con

minimus Candéze, 182

minor Candéze, 182

minusculus Candéze, 1874, Lacon, 39

minusculus Candéze, 1874, Meristhus, 236

minutus Candéze, 22, 39, 95

minutus Schwarz, 1898, 182

minutus Schwarz, 1903, 48

mirificus Candéze, 106

miser Schwarz, 1905) : 281, Lobotarsus

misticius Mignot, 53, 55, 82

mixta Candéze, 18780 : ii (6), Adelocera

mixtus Candéze, 183

miyakei Ohira, 183

miyakoensis Ohira, 183

miyamoti Nakane & Kishii, 183

mjobergi Elston, 184

mocquerysi Fleutiaux, 1899, 237

mocquerysi Fleutiaux, 1902, 106

mocquerysi Fleutiaux, 1932, 122, 184

mocqueryst Fleutiaux, 1934, 122

modestus Boisduval, 56, 71, 136

modestus Candéze, 184

modestus Fleutiaux, 1889 : 139, Lacon

modestus Schwarz, 255

moerens Germar, 114, 240

moestus Candéze, 253

molestus Candéze, 184

molitor Candéze, 185

monachus Candéze, 1882 : 12, Lacon

Monocyrton Golbach, 53, 55, 268

monstrvosum Pic, 13, 16, 26

monstruosus Pic, 26

montanus Miwa, 1929 : 229, Lacon

monticola Candéze, 72

morosus Candéze, 1895a : 48, Lobotarsus

moseri Schwarz, 255

mouhoti Candéze, 1874, 35

mouhoti Candéze, 1888, pars, 31, pars, 35

mouhoti Candéze, 1891, pars, 31, pars, 32

mouhoti Fleutiaux, not Candéze, 30

mozambicana Fleutiaux, 185

mucorea LeConte, 186

mucoreus Candéze, 185, 202

mucronatus Candéze, 255

multiforis Candéze, 185

multipunctatus Elston, 1927 : 358, Lacon

murinus Linnaeus, 20, 54, 113, 114, I15, 116,
118, 119, 186, 209, 240

INDEX 395

murrayensis Blackburn, 186
muscerda Candéze, 115, 187
muscosus Candéze, 187

musculus Candéze, 187

mustellinus Germar, 18, 20, 54, 188
muticus Fleutiaux, 174

muticus Helf. nomen nudum, 189
muticus Herbst, 20, 53, 188
myamoti Nakane & Kishii, 113, 189
Myrmodes Candéze, 113, 115, 268
mysticus Candéze, 189

nagaoi Ohira, 189

nanus Boheman, 40, 280

nebulosus Candéze, 1857, 15, 40
nebulosus Candéze, 1874, 99, 208
nebulosus Fleutiaux, 1889, 1895, 45
nebulosus Razumowskii, 1789 : 161, Elater
nebulosus Schwarz, 208

Neolacon Miwa, 113, 116, 268
nepalensis Candéze, 140

nesiotes Van Zwaluwenburg

ngokoensis nom.n., 190

niger Schwarz, 1902, 41

niger Scharz, 1905, 190

nigrescens MacLeay, 1888 : 1234, Lacon
nigrita Candéze, 190

nigritulus Candéze, 237

nigvoplagiatus Blanchard, 71
niponensis Lewis, 232, 234

nitidus Candéze, 13, 41

nitidus Fleutiaux, 256

nivalis Fleutiaux, 190

nobilis Fall, 53, 72

nodicollis Candéze, 191, 278

nodieri Fleutiaux, 191

nodifer Klug, 20, 191

normandi Buysson, 112

notodonta Latreille, 114, 240, 256, 257
novus Fleutiaux, 73

nubilus Candéze, 1857, Dilobitarsus, 99
nubilus Candéze, 1857, Tilotarsus, 191

oberndorferi Schwarz,
192

oberndorferi Schwarz, 19026 : 202, Pericus, 42

obesus Candéze, 15, 42

oblongus Fleutiaux, 42

obscura Fleutiaux 1934, 192

obscura Szombathy, 215

obscuripennis Fleutiaux, 63

obscurus Fleutiaux, 1934, 192

1902b : 197, Lacon,

obscurus Fleutiaux, 1935, 120

obscurus Fleutiaux, 1940, 73

obtectus Say, 87, 89, 276, 295

occidentalis Candéze, 5, 14, 17, 19, 20, 21, 43

oceanicus Van Zwaluwenburg, 237

occelatus Candéze, 257

Ocneus Candéze, 52, 54, 268

octavus Candéze, 115, 192

Octocryptus Fleutiaux 4, 6 (key), 118, 266,
269

okinawana Ohira, 1967): 101,
Adelocera amamiensis (Ohira)

olcesii Fairmaire, 257

oliveri Candéze, 73

oliviert Schwarz, 73

olsoufieffii Fleutiaux, 64, 73

opacus Candéze, 1878, 257

opacus Candéze, 1900 : 78, Tilotarsus

Opatelus Candéze, 6 (fig.) 8 (key), 18, 19,
91, 268

operculatus Say, nomen nudum, 88

ordinatus Fleutiaux, 112

orientalis Fleutiaux, 73

orientalis Hope, 192, 273

ornatellus Candéze, 193

ornaticollis Castelnau, 1840 : 248, Agrypnus

ornatulus Candéze, 1897 : 11, Meristhus, 232

ornatus Candéze, 193

orthoderus Elston, 1924 :

oshimanus Ohira, 193

ovalis Germar, 5, 13, 14, 15, 16, 17, 20, 21, 54

subsp. of

204, Lacon

pacificus Candéze, 258
paenulatus Boheman, 115, 193, 280
paeninsularis Fleutiaux, 42
paleatus Champion, 117, 118, 194
paleolatus Candéze, 92

palliatus Candéze, 194

palliatus Latreille, 74
palliditarsus Candéze, 194
pallidus Candéze, 195

palpalis Candéze, 195
pantherinus Fleutiaux, 195
Payralacon Reitter, 113, 116, 268
paralella Lewis, 75

paralellicollis Candéze, 1857, 195
paralellicollis Candéze, 1889, 258
paralellus Candéze, 196
paralellus Schwarz, 106

parcus Boheman, 42

parcus Schwarz, 196

pardalinus Candéze, 1882 : 9, Lacon
parviceps Gyllenhal, 196

306 C. M. F. von HAYEK

parviceps Schoenherr, 20, 54, 196
parvulus MacLeay, 1888 : 1239, Lacon
parvus Fleutiaux, 196

pauliani Fleutiaux, 1941d : 38, Lobotarsus
pauliani Girard, 197

pauliani Mouchet, 106

pauper Candéze, 197

pecirkai Jagemann, 10944 : 333, variety of

schwarzi Jagemann, 1944 : 333, Lacon
pectinata Candéze, 75
pectinicornis Champion, 75
pectoralis Fairmaire, 75
pellinaria Golbach, 76
pendleburyi Fleutiaux, 94, 99
pennata Fabricius, 14, 62, 270
Pericus Candéze, 13, 15, 268
peringueyi Candéze, 257, 258
permucronatus Schwarz, 258
perplexus Elston, 197
perraudieri Fleutiaux, 1889, 237
perraudieri Fleutiaux, 1927, 45
perviert Fairmaire, 107
persicus Candéze, 259
pescadoriensis Miwa, 241, 259
petiginosus Germar, 99
petitti Dejean, nomen nudum, 229
philippinus Fleutiaux, 75
picta Fleutiaux, 75
pictilis Schwarz, 197
pictipennis Candéze, 197
pictiventris Candéze, 198
pictus Candéze, 1978) : LIII (6), Lacon
piger Candéze, 198
pinguis Candéze, 198
pipitzi Candéze, 199
pistorius Candéze, 199, 270
pistrinarius Candéze, 237
plagiatus Candéze, 199
planatus Candéze, 199
planus Candéze, 45, 109
pleureticus Candéze, 200
plumatus Fleutiaux, 26
politus Candéze, 259
pollinaria Candéze, 76
ponderatus Candéze, 259
porcatus Linnaeus, 78
porcinus Candéze, 200
porosus Klug, 200
porrecta Fleutiaux, 200
porriginosus Candéze, 118, 200
praelongus Elston, 201
praetermissus Candéze, 201
praetexta Fleutiaux, 201
pretoriensis Cobos, 113, 115, 193

principatus Schwarz, 1905a : 259, Lacon
princeps Candéze, 201

procellosus Candéze, 201

productus Elston, 1924 : 203, Lacon
profusa Candéze, 88

Prolacon Fleutiaux, 13, 15, 268
Prolobotaysus nomen nudum, 156
propinquus Candéze, 151

Protelater Sharp, 5, 101

proxima Fleutiaux, 202

proximus Fleutiaux, 241, 259
pruinosus Fairemaire, 1849 : 35, Agrypnus
Psephus, I1

Pseudolacon Blackburn, 113, 115, 268
Psilonicus, 86

puber Candéze, 12, 260

pubescens Candéze, 260

pulchellus Schwarz, 106

pulcher Schwarz, 106

pulvereus Candéze, 185, 202
pulverulentus Herbst, 78
pulverulentus Fabricius, 68, 78
pulverulentus Panzer, 76

pumilis Candéze, 260

punctatissimus Elston, 202

punctatus Candéze, 253

punctatus Herbst, 52, 54, 55, 56, 75, 240
punctipennis Candéze, 118, 202
pupillus Candéze, 203

pusillus Schwarz, 50

Pyrganus Golbach, 113, 117, 269
pygmaeus Baudi, 12, 19, 20, 21, 45
pyrsolepis LeConte, 53, 78

quadricollis Fairmaire, 203

quadricollis Fleutiaux, 112

quadriguttatus Candéze, 45

quadrilineatus Hope, 272

quadrimaculatus Fabricius, 114

quadrinodatus Lewis, 53, 55, 78

quadrinotatus Schwarz, 203

quadyinotulus Schenkling, 203

quadripicturatus Schwarz, 1891 : I5I,
Lacon

quadripunctatus Candéze, 231, 234, 235

quadrituberculatus Candéze, 99

quedenfeldti nom. n., 204

queyca Reitter, 78

quercea Herbst, 52, 78

quercinus Gmelin, 78

quercus Fuesley, 79

quercus Seidlitz, 78

INDEX 397

radula Candéze, 266, 267

raffrayi Candéze, 104, 107

ramitarsus Ca déze, 7 (fig.), 239, 240
ramatasenseni Miwa, 1934 : 180, Adelocera
ramusculus Candéze, 107

rectangularis Say, 113, 118, 146, 204
rectangularis Schwarz, 1903b : 378, Lacon
recticollis Elston, 204

recticollis Fleutiaux, 79

reductus Candéze, 1878, 158, 205
veductus Candéze, 1882, 158

veedi Candéze 92, 93

renardi Candéze, 261

resectus Candéze, 261

reticulatus Elston, 205

Rhaciaspis Arnett, 231, 269

riehlii Candéze, 52

ripario Germar, 18

riparius Fabricius, 18

Rismethus Fleutiaux 7 (key), 235, 269
vitsemae Candéze, 37

rivularis Gyllenhal, 18

robusta Fleutiaux, 79

robustus Fleutiaux, 261

robustus Schwarz, 1903b : 377, Lacon
rohanchaboti Fleutiaux, 205

rorulenta LeConte, 79

ruber Perty, 80, 114 (and see rubra Perty)
rubescens MacLeay, 1888 : 1234, Lacon
rubicundulus MacLeay, 1888 : 1237, Lacon
rubicundus Candéze, 45

rubida Schwarz, 80

rubiginosus Candéze, 1865, 261
rubiginosus Candéze, 1882, 205

rubra Perty, 80

rubripennis Fleutiaux, 80

tufipes Candéze, 262

tufipes DeGeer, 1774 : 150, Elater
rufopictus MacLeay, 1888 : 1235, Lacon
rufulus Elston, 1927 : 361, Lacon

rufus Blackburn, 113, 115, 206

rufus Candéze, 262

rugatus Candéze, 206

Tugosa Fleutiaux, 206

rusticus Candéze, 206

Sabikikorius Nakane & Kishii, 113, 117, 269
sachalinensis Miwa, 80

Sagojo Ohira, 113, 117, 269

Sagojyo Kishii, 113, 117, 269

sahlbergi Schwarz, 89

| sakaguchii Miwa, 206

sakashimanus Ohira, 132

sallei LeConte, 262

salvazai Fleutiaux, 81

samburensis Fleutiaux, 207
sanguinea Fleutiaux, 54, 71
sanguineus Candéze, 52, 54, 81
sanguinolentus Candéze, 41, 270
santali Fleutiaux, 46

sauteri Ohira, 1970b : 208, Adelocera
scaber Candéze, 207

scabrosus Fleutiaux, 234
Scaphoderus Candéze, 8 (key), 51, 268
scarrosus Candéze, 117, 207

Scelisus Candéze, 52, 54, 268
schonfeldti Candéze, 46

schotti LeConte, 240, 243, 263
schwaneri Candéze, 207

schwarzi Jagemann, 1944 : 333, Lacon
schwarzi Schenkling, 208

scobinula Candéze, 231, 232, 235, 238
scobinula Lewis, 239

scopulosus Elston, 208

scopus Schwazrz, 208

scortecci Binaghi, 263

scrofa Candéze, 208, 277

scrobinula Candéze, 231

scrofilia Ohira, 209

sculptus Candéze, 209

scutellaris Candéze, 209

scutellatus Candéze, 165, 209
scutellatus Fairmaire, 165
semicribrosus Fairmaire, 264
semistriatus Schwarz, 264
semivestitus Elston, 210

senegalensis Castelnau, 240, 256
senegalensis Dejean nomen nudum, 114
senilis Germar, 14, 65

senilis Péringuey, 1892 : 44, Lacon
septentrionalis Fleutiaux, 210
sericans Candéze, 1857 : 112, Lacon
serricollis Candéze, 210

serricornis Quedenfeldt, 47

serrula Candéze, 210

setarius Chevrolat, 238

setiger Bates, 118, 163, 211

setosa Candéze, 81

setosus Schwarz, 2II

setulosus Candéze, 211

setulosus Schwarz, 19036 : 378, Lacon
seyrigi Fleutiaux, 211
shirakii Matsumura, IQII :
shirozui Ohira, 13, 47
signatus Candéze, 43
signifer Candéze, 107
silvatica Fleutiaux, 212

146, Lacon

308 €. M. FF: von HAYEK

similis Fleutiaux, 212

simplex Candéze, 212

sinensis Candéze, 116, 212

singularis Fleutiaux, 213

sinuatus Candéze, 118, 213, 278

sjostedti Schwarz, 213

sobrinus Candéze, 1887, 264

sobrinus Candéze, 1895, 108

socius Candéze, 213

soleatus Candéze, 214

sonadaicus Candéze, 253, 254

sovdidus Candéze, 186

soricinus Candéze, 1857, 214

soricinus Candéze, 1882, 265

sparsa Candéze, 81

sparsus Candéze, 47

specularis Van Zwaluwenburg, 1947 : 109,
variety of gracilis Candéze

spinifer Candéze, 214

spissicollis Candéze, 214

spretus Candéze, 1882 : 8, Lacon

spurca Candéze, 54, 82

spurcus Candéze, 1865 : 11, Lacon

squalescens Blackburn, 156

squalescens Fairmaire, 156, 214

squalidus Fairmaire, 1849 : 35, Agrypnus

squalida Fleutiaux, 215

squamea Szombathy, 215

squameus Candéze, 233

squamiger Champion, 235, 239

stictus Candéze, 215

stigmosus Elston, 118, 215

strangulata Fleutiaux, 82

striatus Fleutiaux, a variety of extractus
Fleutiaux, 265

stricticollis Fairmaire, 216

suarezl nom. n., 216

subaurorata Schwarz, 1904 : 46, Adelocera

subcarinulatus Schwarz, 216

subcervinus Fleutiaux, 217

subcompactus Elston, 1927 : 356, Lacon

subcostata Candéze, 55, 82

subcostatus Candéze, 48

subcylindricus Schwarz, 217

submarmoratus Elston, 217

subocellatus Candéze, 217

suboculatus Candéze, 218

subpectinata Schwarz, 52, 54, 83

subsericeus Candéze, 218

subserratus Quedenfeldt, 218

substriata Fleutiaux, 218

substriatus Candéze, 265

subsulcatus Candéze, 100

subtilis MacLeay, 1888 : 1236, Lacon

subtuberculata Fleutiaux, 173
subtuberculatus Schwarz, 173, 218
succinatus Candéze, 48, 273

succinea Hope, 273

suillus Candéze, 219

sulcicollis Schwarz, 216, 219

sulcicollis Solier, 93, 94, 100

sulcifrons Candéze, 1893c : 171, Lacon
Sulcilacon Fleutiaux, 52, 54, 268
Sulcimerus Fleutiaux, 7 (key), 231, 269

tabaci Fleutiaux, 48

tabularius Candéze, 1892c¢ : 483, Lacon

taciturnus Candéze, 141, 219

tactus Candéze, 219

taeniata Panzer, 1795 : 236, Elater

tanganus nom. n., 48

taiwanus Miwa, 220

tectus Fleutiaux, 220

tellini Fleutiaux, 148, 220

tenebrionoides Candéze, 83

tenebryioides Candéze, 83

tenuicollis Kishii, 1961 : 15, aberrant form of
binodulus Motschulsky

terminata Fleutiaux, 220

tervenus Germar, 20, 54, 144, 188, 196, 274

tessellata Candéze, 1893) : 6, Adelocera

tessellatus Candéze, 100

testaceus Fleutiaux, 23, 49

testaceus Schwarz, 220

tetraspilota Hope, 273

texanus Horn, 232, 238

thomasi nom. n., 221

thibetanus Reitter, 221

thoracicus Scopoli, 1763 : 92, Elater

tigrinus Fleutiaux, 221

Tilotarsus Candéze, 113, 115, 268

tippooi Candéze, 265

tomentosus Fabricius, 114, 240, 266

tonkinensis Fleutiaux, 1927, Adeloceta, 221

tonkinensis Fleutiaux, 1927, Agraeus, 35

tonkinensis Fleutiaux, 1927, Alaotypus, 83

tonkinensis Jagemann, 1944 : 334, Compso-
lacon

torrefactus Candéze, 222

torresi Candéze, 222

tostus Candéze, 222, 271

Trachylacon Motschulsky, 13, 15, 268

transversicollis Fleutiaux, 222

transversus Candéze, 222

triangularis Schwarz, 1903@ : 362, Lobo-
tarsus

Triers Candéze 7 (fig., key), 239, 269

INDEX 309

trifasciatus Candéze, 1865, 38

trifasciatus Candéze, 1873, 28

tripartitus Candéze, 233

tripartitus Fleutiaux, 49

tristis Blanchard, 1853 : 86, Lacon

tristis Kraatz, 65, 67

Trochylacon Motschulsky, 33

troglodytes Boheman, 40, 280

tropicus Hope, 11, 260

truncatus Herbst, 115, 118, 172, 182, 223

truquii Candéze, 223

tuberculatas Latreille, 93, 95

tuberculatus Candéze, 224

tuberculipennis Miwa, 1929 : 230, Lacon

tumens Candéze, 49, 277

tumens Candéze sensu Fleutiaux, 25

tumidicollis Candéze, 228

tumidipennis Candéze, 50

tumulosa Candéze, 84

turbatus Candéze, 224

turbidus Germar, 20, 53, 54, 127, 130, 188,
194, 224, 274

turgescens Candéze, 50

turkistanicus Schwarz, 225

tuspanensis Candéze, 113, 117, 225

Tylotarsus Germar, 113, 115, 116, 268

typicus Kishii, 1961 : 14, aberrant form of
binodulus Motschulsky

undulatus Schwarz, 108

unicolor Candéze, 84

unicolor Hope, 113, 115, 225
unicus Fleutiaux, 226

uraiensis Miwa, 1929 : 231, Lacon
ursulus Candéze, 118, 226

validus Elston, 1924 : 203, Lacon
vandepolli Candéze, 204, 226
variabilis Candéze, 13, 205, 226, 227
variatus Candéze, 227

variatus Schenkling, 227

C. M. F. von Hayek, B.Sc.
Department of Entomology

British MusEum (NATURAL History)
CROMWELL RoapD

Lonpon SW7 5BD

variegatus Fleutiaux, 1902, 108

variegatus Schwarz sensu Fleutiaux, 1918,
1927, 33

variegatus Motschulsky, 1854 : 45, Lacon

variegatus Schwarz, 1902, 50

variegatus Schwarz, 1906, 227

variolus Candéze, 228

varius Fabricius, 53

varius Oliver, 14, 52, 53, 78, 79, 114, 271

versicolor Fleutiaux, 228

vestitus Klug, 20, 54, 114, 118, 228

veternosus Elston, 1930 : 22, nomen nudum,
see also Neboiss, 1956 : 14

vetusta Walker, 88

vicina Candéze, 84

vicinus Candéze, 108

vicinus Fleutiaux, 229

victoriae Candéze, 229

viettei Girard, 1970, Archontas, 229

viettei Girard, 1970, Lacon, 84

vivvoratus Candéze, 171

villiersi Cobos, 112

viridis Champion, 85

vitalisi Fleutiaux, 1918, Adelocera, 54, 85

vitalisi Fleutiaux, 1918, Lacon, 229

vitticollis Fairmaire & Germain, 86, 94, 95,
97, 100

wallacei Candéze, 1874, Adelocera, 85
wallacei Candéze, 1874, Lacon, 124, 230
weisei Schwarz, 1898) : 183, Hemicleus, 112
weisei Schwarz, 1898) : 185, Lacon, 51

yayeyamana Miwa, 1934 : 246, Adelocera
yilgarnensis Blackburn, 144

yunnanus Fleutiaux, 85

yuppe Kishii, 113, 117, 230

Zalepia Arnett, 53, 55, 268
zanzibaricus nom. n., 230
zietzi Blackburn, 230

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177:

18 plates, 270 text-figures. August, 1965. £4.20.

. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,

Termitidae) from the Ethiopian Region. Pp. 172 : 500 text-figures. September,
1965. £3.25.

. OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-

philidae. Pp. 129 : 328 text-figures. May, 1966. 3.

. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family

Coccidae (Homoptera : Coccoidea). Pp. 168 : 43 text-figures. January, 1967.
£3.15.

. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the

world species of Cleora (Lepidoptera : Geometridae). Pp. 119: 14 plates, 146
text-figures, 9 maps. February, 1967. 3.50.

. HemminG, A. F. The Generic Names of the Butterflies and their type-species

(Lepidoptera : Rhopalocera). Pp. 509. £8.50.

. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera :

Rhopalocera). Pp. 322 : 348 text-figures. August, 1967. £8.

. Mounp, L.A. Areview of R.S. Bagnall’s Thysanoptera Collections. Pp. 172:

82 text-figures. May, 1968. {4.

. Watson, A. The Taxonomy of the Drepaninae represented in China, with

an account of their world distribution. Pp. 151: 14 plates, 293 text-figures.
November, 1968. {5.

. AFIFI, S. A. Morphology and Taxonomy of Adult Males of the families

Pseudococcidae and Eriococcidae (Homoptera : Coccoidea). Pp. 210: 52 text-
figures. December, 1968. {5.

. CROSSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and

its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75.

. ExroT, J.N. Ananalysis of the Eurasian and Australian Neptini (Lepidoptera :

Nymphalidae). Pp. 155 :3 plates, ror text-figures. September, 1969. {4.

. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe

(Hymenoptera : Chalcidoidea). Pp. 908 : 686 text-figures. November, 1969.
£19.

. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198:

68 plates, 15 text-figures. October, 1971. 12.

. SANDS, W. A. The Soldierless Termites of Africa (Isoptera : Termitidae).

Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90.

. CROSSKEY, R. W. A revisionary classification of the Rutiliini (Diptera;

Tachinidae), with keys to described species. Pp. 167:109 text-figures.
February, 1973. £6.50.

PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND

eS

ne a

im.

A CONSPECTUS
OF THE TACHINIDAE (DIPTER.
OF AUSTRALIA, INCLUDING© KEYS
TO THE SUPRASPECIFIC TAXA”,
AND TAXONOMIC AND HOS?”
CATALOGUES

R. W. CROSSKEY
:
|
|

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ee LOGY Supplement 21
LONDON : 1973

:

A CONSPECTUS OF THE TACHINIDAE (DIPTERA)
OF AUSTRALIA, INCLUDING KEYS TO THE
SUPRASPECIFIC TAXA AND TAXONOMIC,

AND HOST CATALOGUES ee, :)

XW,

Se” 7 niS\
s AA

BY
ROGER WARD CROSSKEY

Pp. 1-221; 95 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ENTOMOLOGY Supplement 21
LONDON : 1973

THE BULLETIN OF THE. BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be
completed within one calendar year.

In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.

This paper is Supplement 21 of the Entomological
series. The abbreviated titles of periodicals cited
follow those of the World List of Scientific Periodicals.

World List abbreviation
Bull. Br. Mus. nat. Hist. (Ent.) Suppl.

© Trustees of the British Museum (Natural History), 1973

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued 7 December, 1973 Price £9.55

meCONSPECTUS OF THE TACHINIDAE (DIPTERA)

PE AUSTRALIA, INCLUDING KEYS TO THE
SUPRASPECIFIC TAXA AND TAXONOMIC

aN HOST CA TREOGUES

By R. W. CROSSKEY

CONTENTS

SYNOPSIS
PREAMBLE

Part I — KEyS TO THE SUPRASPECIFIC TAXA OF AUSTRALIAN TACHINIDAE
Introduction :
Material and methods
An annotated glossary of characters and terms used i in the —
Key to Australian subfamilies of Tachinidae
Subfamily Phasiinae with keys to the tribes and genera
Subfamily Proseninae (Dexiinae) with keys to the tribes and genera
Subfamily Tachininae with keys to the tribes and genera .
Subfamily Goniinae with keys to the tribes and genera

Part II — A TAXONOMIC CATALOGUE OF THE AUSTRALIAN TACHINIDAE

Introduction

Explanatory information on the Catalogue format :

Synopsis of the catalogue arrangement of tribes and genera

The taxonomic catalogue

Summary of nomenclatural changes established i in the catalogue

Lectotype designations .

Summary of Australian nominal species “for which types are lost or
missing :

Part III — A HOST CATALOGUE FOR THE AUSTRALIAN TACHINIDAE .

Introduction ; :
A synopsis of the host-relations of Australian Tachinidae :
Parasite-host list .
Host-parasite list .

ACKNOWLEDGEMENTS

REFERENCES

ILLUSTRATIONS

APPENDIX

INDEX TO FAMILY-GROUP NAMES.

INDEX TO GENUS-GROUP NAMES .

INDEX TO SPECIES-GROUP NAMES

Page

ON ADO +P

26
29
41
50
74

Ioo

100
100
105
109

157
160

165

166

166
169
171
178
185
186
194
209
210
211
215

4 R. W. CROSSKE ¥

SYNOPSIS

A detailed conspectus is provided of the Australian Tachinidae, a group that has hitherto
lacked any taxonomic synthesis. Original keys are given to the family-group taxa and to
the genera and subgenera so far recognized in the fauna; many undescribed forms are known
and the preliminary nature of the keys is therefore emphasized. Preliminary characteriza-
tions are given for family-group taxa to aid towards formalized diagnoses on a world basis.
A complete systematic catalogue is given of all described Australian Tachinidae, based upon
an examination of almost all extant primary types, and the known host relations are shown
by up-to-date host-parasite and parasite-host lists. Nomenclatural changes established in
the taxonomic catalogue are summarized and include 14 new genus-group synonyms, 26 new
species-group synonyms, 83 new generic combinations, and two new names for preoccupied
homonyms; 12 lectotypes are newly designated. An annotated glossary is included of the
terms used in the taxonomy of adult Tachinidae, and figures are given to illustrate the glossary
and the keys.

PREAMBLE

AUSTRALIA possesses a rich and varied tachinid fauna, most of which is still
undescribed. Some 420 named species are recognized at present, but it is obvious
from an examination of unidentified material in the collections at Canberra and
London that this number represents only a small proportion of the species that
will ultimately be recognized: it is probable that when fully worked out the
Australian Tachinidae will muster some 1500-2000 species.

This fauna, though not enormous by the standards of some insect families,
presents considerable taxonomic difficulties (Colless & McAlpine, 1970) and
it will certainly be many years before any comprehensive monographic treatment
can be prepared and the multifarious new genera and species adequately described
and named. In the meantime the practical identification of Australian Tachinidae
is bound to present a problem for which the services of a specialist in the group
are likely to be required, if only to distinguish rapidly the known from the unknown
elements in the fauna. At present there is no full-time Australian specialist
working on the group, despite the fact that there is a very real need for taxonomic
studies to be undertaken on the Australian tachinids because of their role as
parasites of other insects and their potential importance as biological control
agents. Australia suffers from many serious insect pests of economic significance
(such as the chafer-grubs of sugar-cane and the Eucalyptus defoliators), all of
which are attacked by tachinid parasites that are assumed to play an important
part in naturally regulating the populations of their hosts; few attempts have
yet been made, however, to exploit the Tachinidae of Australia for biological
control purposes.

It will be an essential prerequisite for future investigations into the value of
Australian tachinids as biological control agents that the flies themselves can be
rapidly and accurately identified. At the moment only a small fraction of the
fauna can be so named, since the great bulk of species remain unstudied and
undescribed. An enormous untouched taxonomic field, with a direct bearing on
the practical control of insect pests, awaits the Australian student willing to take
up the systematics of the Australasian Tachinidae as a whole (attention cannot

TACHINIDAE OF AUSTRALIA 5

be paid solely to continental Australia as much of the fauna is held in common
with that of New Guinea and the Pacific islands and in some instances with areas
still further afield).

Up to now Australian dipterists have been deterred from working on their
local tachinid fauna by the practical difficulties of knowing where to start in the
absence of any comprehensive revisionary works and scarcely any keys, and by
the fact that so many of the type-specimens are housed in collections outside
Australia. For some time there has been a need for a synthesis of existing
taxonomic knowledge which will provide a foundation upon which future work
can be developed, and the object of the work here presented has been to provide
a synthesis of this kind. The work has been based on a study of the Australian
Tachinidae carried out at intervals over the last ten years, and its aims are to
provide: (a) a classification of described forms and a classificatory framework
into which new forms can be fitted, with whatever modifications may be necessary,
as they are described; (b) preliminary characterizations of the subfamilies and
tribes recognized in the fauna and keys to family-group taxa; (c) identification keys
to the described genera and subgenera; (d) a taxonomic catalogue, based upon an
examination of all available primary types and geographically annotated; (e) a
catalogue of known hosts; and (f) an illustrated glossary of the terms used in the
taxonomy of adult Tachinidae that will aid the would-be student in acquiring a
knowledge of the group. It has not been practical at this stage, when many genera
remain in need of complete revision, to provide keys: to species and descriptions
of species, and it should be noted that some of the species names listed in the
catalogue may prove to be synonyms of other names when their genera are
studied in detail.

Finally in this preamble it might be useful to comment briefly on the apparent
affinities and zoogeographical relationships of the Australian Tachinidae. In the
main the fauna consists of endemic genera and species occurring principally in the
eastern and southern parts of the Australian continent and in Tasmania, but in
northern Queensland and in the Northern Territory this essentially Australian
fauna is supplemented by many Oriental genera and species that occur widely
throughout South-East Asia and spread eastwards into Melanesia and northern
Australia. There is thus a large shared element in the fauna between New Guinea
and Queensland of forms that probably reached Australia by immigration from
the north and west. In addition to this, however, there has perhaps been a
contrary movement of characteristically Australian forms northwards into New
Guinea (for recent collecting has now shown the presence of such typically
‘Australian’ genera as Amphibolia, Chaetophthalmus and Tritaxys in the central
New Guinea highlands), unless the common elements between upland New

| Guinea and upland New South Wales are separated remnants from a formerly

widespread distribution. A few species occurring in Australia are widespread

_ throughout the Old World, and it seems likely that critical future work will show
_ the presence in Australia of species having a circum-Indian Ocean distribution

from eastern Africa through peninsular India and on to Western Australia or

| Queensland: some tachinid parasites, such as Carcelia species attacking Heliothis

6 R. W. CROSSKEY

cotton bollworms, masquerade under different names in different zoogeographical
regions (just as their hosts often do) when almost certainly only a single species
is involved. Between Australia and New Zealand there is almost no relationship
at all in the tachinid fauna, that of New Zealand being a baffling and peculiar
fauna very different from that of the rest of the Old World; the only notable point
of resemblance between the Australian and New Zealand faunas is in the Phasiini,
where the New Zealand ‘genus’ Campbellia suggests derivation from the Australian
part of the Alophora (Mormonomyia) complex. Little can be said about any
possible relationship between the Australian Tachinidae and those of South
America beyond commenting that there is some resemblance in facies between
forms in Tasmania (and nearby parts of the Australian mainland) and some forms
in the southern Neotropical fauna; the significance of this resemblance is not clear.

All the main subfamilies and tribes of Tachinidae are represented in Australia,
but the area is remarkable for the rich development of the Proseninae (=Dexiinae)
and in particular of the Rutiliini (Crosskey, 1973). The abundance of forms in this
tribe appears to be closely correlated with the richness in Australia of the chafer
fauna (Scarabaeidae : Melolonthinae), which provides the hosts for these tachinids.

PART I—KEYS TO THE SUPRASPECIFIC TAXA OF AUSTRALIAN TACHINIDASB

INTRODUCTION

It is certain that the several hundred species of Tachinidae known to occur
in Australia represent only a small proportion of the fauna that will ultimately
be discovered and (presumably) named. It is therefore premature, in a sense,
to attempt to provide keys to the supraspecific taxa, especially when experience
shows that even with a well-known fauna like the Tachinidae of Britain it is
difficult to construct really satisfactory keys that anyone but a specialist can use
reliably. Yet in attempting to acquire a knowledge of a large and complex insect
fauna like the Tachinidae of Australia the potential student is in need of some
keys that at least begin to organize the mass of data available and to show how the
many described genera can be differentiated, and how the higher taxa to which
they belong can best be recognized.

Scarcely any keys to the Australian fauna have up to now been available.
Malloch (various papers) published small piecemeal keys to place some of his
newly described taxa among their relatives, and one or two longer keys for the
recognition of artificial groups of genera, but these have been of very limited use
and are now outmoded by changes in generic concepts that have taken place
during the past thirty years or so. Apart from these, the only keys published
to any Australian supraspecific taxa have been restricted to the Rutiliini
(Paramonov, 1968; Crosskey, 1973).

None of these pre-existing keys is of any use in providing the would-be student
of the Australian fauna with an over-all system of keys that will provide a means
of identifying the many tribes and genera to be found on the Australian continent.
The object of these keys here presented is to provide just such a system, so that

TACHINIDAE OF AUSTRALIA 7

the worker interested in taking up this group has some means at hand for
beginning on a study of the Tachinidae without necessarily having immediate
access to a large museum collection: to aid the student as much as possible a
detailed glossary is given of all the main terms used in the keys.

The Tachinidae is a taxonomically very difficult family, as Colless & McAlpine
(1970) have emphasized in their account of the Australian Diptera, and it is
notoriously difficult to make keys that are free from all possibility of error when
identifying specimens. Specialists on the family frequently find difficulty in
using keys, and not infrequently make errors of identification in spite of their
knowledge of the family; these points are emphasized, so that the beginner on
the group shall not feel too discouraged when keys appear to fail (as they will
occasionally, since much of the fauna remains unknown) or when the specimen
that ran out so convincingly to a certain name proves to be something quite
different.

MATERIAL AND METHODS

The keys and diagnostic matter are mainly based on a study of material in the
British Museum (Natural History), London, together with a study of types
(especially those of the type-species of Australian genera) from the collections in
Berlin, Canberra, Eberswalde, Ottawa, Paris, Vienna and Washington. The
BMNH collection is the largest and most representative of world Tachinidae, and,
except for the Australian National Insect Collection, is richer in material from
Australia than other collections.

With few exceptions the early stages of Australian Tachinidae remain completely
unknown and the keys are, perforce, based only on adult characters. For
describing these the following conventions and abbreviations are used.

In describing the positions of leg setae the convention is followed of imagining
the leg to be extended at a right-angle to the longitudinal axis of the fly, when:

a anterior p _ posterior

ad anterodorsal pd_ posterodorsal
av anteroventral pv posteroventral
d dorsal v ventral

A tibial seta indicated by any of these letters is on the shaft of the tibia and not at its
end unless otherwise specified.

The abbreviations used for thoracic setae are:

acy acrostichal pra pre-alar

dc dorsocentral prst acy presutural acrostichal
wa intra-alar prst dc presutural dorsocentral
ph posthumeral prst ia presutural intra-alar
post acy postsutural acrostichal sa supra-alar

post dc postsutural dorsocentral stl sternopleural

post ia postsutural intra-alar

8 R. W. CROSSKEY

Abdominal tergites are indicated by the letter T followed by the appropriate
number; the composite first apparent tergite is Tr + 2, the usual last visible
tergite T5. Abdominal sternites are indicated by the letters St suffixed as for
tergites (only the male St5 generally requires citation).

Parts of the male hypopygium are infrequently cited in the keys, but the
terminology used by Colless & McAlpine (1970) is used throughout the paper
whenever genital features are mentioned.

All keys are regularly dichotomous. The keys include names of a few genera
that are not yet positively known from Australia but seem likely to be found
there: in such cases the names are printed in non-bold type. Where a generic
name is included in square brackets in the keys it indicates that the tribal position
of the genus concerned is uncertain, but that the genus is included in a different
tribe in the formal classification adopted from the one to which the key relates.

Figures have all been drawn personally and attempt to show only the essentials
required for identification (needless shading and vestiture have been omitted).
An attempt has been made to illustrate basic patterns of chaetotaxy (see
Text-figs 4, 7 & 54-63) on the thorax by omitting the bristles themselves and
indicating their distribution just by the ‘pore-patterns’ of their insertions.
Such a method of illustrating chaetotaxy seems hardly to have been used at all
in tachinid taxonomy but provides a useful visual aid for recollecting the most
fundamental and frequently recurring patterns. It must be emphasized that the
circles indicating the bristle pores are exaggerated in size relative to the sclerites,
but that different sized circles are used to show (approximately) the relative sizes
of the bristles to each other. Broken lines between circles indicate the serially
arranged setae that have the same composite terminology.

AN ANNOTATED GLOSSARY OF CHARACTERS AND TERMS USED IN THE KEYS

The glossary here given summarizes the terminology used in the keys, so as
to make these as comprehensible as possible to the non-specialist (including in
particular the Australian student who might wish to take up tachinid taxonomy).
Hardly any of the recent works —and very few old works — contain any glossary
of the terms habitually used by taxonomists working on the Tachinidae, and the
glossary here presented ought (it is hoped) to be of benefit to my specialist
colleagues in so far as it attempts to define the external adult characters most often
used in supraspecific taxonomy and to correlate the various synonymic terms
most commonly used by different authors.

The terminology adopted is that which appears to be the most universally
accepted, and most readily comprehended, by specialists. It is, however, essentially
a taxonomist’s vocabulary, and some of the terms are at variance with those
favoured by the morphologist. This point is specially germane when dealing
with the Australian fauna, as Colless & McAlpine (1970) in their work on the
Australian Diptera have adopted a strongly morphological line for their structural
terminology, and there are therefore some discrepancies between the taxonomic
terminology and that of Colless & McAlpine; in particular this affects the names

TACHINIDAE OF AUSTRALIA 9

of parts of the thorax and of certain wing veins. Some of the terminology used
by Colless & McAlpine is helpful for the taxonomist, especially their terms
pleurotergite and mediotergite for parts of the thorax that have not had satisfactory
names in the taxonomic literature, and these terms are here adopted (as
pleurotergite, for example, is a much handier name than the usual swpra-spiracular
convexity of the taxonomist); on the other hand it is disadvantageous in taxonomy
to have to speak of the posterior pronota and in this case I prefer the time-honoured
taxonomic term humeral calli. In regard to the nomenclature of wing veins I
continue to follow the standard work on Tachinidae by Mesnil (1944) in preference
to the venational notation found in most of the more generalized works on Diptera
(including that of Colless & McAlpine); the main difference between these systems
that is of practical importance concerns the so-called fifth vein which Mesnil
(here followed) calls Cu, and which is known as M,,, in other works (e.g. Colless
& McAlpine, 1970).

The existence of many alternative names for different structures has made it
necessary to record the more important or frequent alternatives, and this has
been done in the glossary by entering the alternative name(s) when needed
(italicized and in parentheses) at the end of each definition. The alternatives
should be helpful in correlating the present work with that of other authors (for
example, by showing that the structures here termed parafrontals are those often
known as orbits). In some instances the German equivalents have been given so
that the English terminology used in keys can be readily associated with the
German terminology used in Mesnil’s very important work on Tachinidae in
Lindner’s Die Fliegen der Palaearktischen Region 64g (which contains keys of
great value on a world basis as well as for the local Palaearctic fauna).

Many of the glossary definitions are accompanied by separately paragraphed
annotations that are designed to show, very briefly, the taxonomic value of the
characters for which the terms stand. In these notes examples are often cited
of particular genera or higher taxa in which a certain character condition occurs:
the examples are drawn from the Australian tachinid fauna, but most of them are
relevant to other zoogeographical regions as well. The annotations eee only
to Tachinidae and must not be read as applicable to other Diptera.

The accompanying Text-figures 1-23 have been specially prepared to illustrate
the characters mentioned in the keys and as an adjunct to the glossary definitions.

abdominal T1-+ 2. The apparent first segment of the abdomen, formed
compositely of fused first and second tergites (loosely, first segment) (Text-figs 91
& 94).
acrostichal setae. The innermost two longitudinal rows of setae on the mesonotum
(Text-fig. 4).
These may be absent or reduced (e.g. in some Phasiini, Minthoini) or
__ represented by only the prescutellar pair.
antennal axis. An imagined horizontal line through the head profile at the level
of the antennal insertions (Text-fig. 14).

10 kh. W CROSSIREY

The height of the axis relative to the eye centre and the head length at this
axis in relation to head length at the epistomal axis can be significant in
describing head form.

apical scutellar setae. The hindmost pair of marginal setae on the scutellum
(except when undeveloped) (Text-figs 5 & 6).

Orientation of these setae (whether horizontal or upright, crossing or
diverging) can be taxonomically important. If the apical setae are unrepresented
then other marginal setae are the hindmost ones in a literal sense.

appendiculate. Provided with an M, appendix (q.v.) (said of vein M/).
appendix. A spur-like vein of varied length that continues almost directly towards
the wing margin from the bend of vein M (designated M,) (Text-fig. 10).

Normally present only when vein M is abruptly angled and sometimes

represented only by a weak vestige or even a mere fold in the wing membrane.
arista. The setiform or style-like part of the antenna arising externally from
the base of the third antennal segment (German: Fiihlerborste) (Text-fig. 1).

Present in all Tachinidae and providing useful taxonomic features in hair
length, extent of thickening, and elongation of the two basal segments.
Typically it is micropubescent (with very inconspicuous hairing that is shorter
than its own diameter) or pubescent (with more conspicuous longer hair that
does not or only slightly exceeds its own diameter), but may be plumose (with
long hairs that greatly exceed in length its own diameter and give the arista
an obvious bushiness). Micropubescent or pubescent in all Phasiinae and
nearly all Goniinae, often plumose in Proseninae and Tachininae (especially
Minthoini).

barette. A small subrectangular area on the pleural region of the thorax
differentiated between the pteropleuron and the hypopleuron (meropleuron)
(Text-fig. 7).

Of minor taxonomic importance in the extent of its hairing. Usually a few
hairs only on anterior part but commonly bare, less often fully haired along
its length (e.g. in most Rutiliini, Winthemia).

basal node of R,,;. The slightly or strongly swollen basal part of wing vein
R,4, near its bifurcation from vein R,,, (Text-fig. 10).

Usually bearing one more more small hairs or setulae on the upper surface
(usually also on lower surface), sometimes totally bare (e.g. in many Phasiini,
a few Blondeliini and Leskiini). Presence of only one very strong setula (e.g.
in Palexorista and allied Sturmiini, many Neaerini and Acemyini) in contrast
to several small hairs is often taxonomically important.

basal scutellar setae. The pair of marginal setae nearest to the scutellar base
(except when, very rarely, undeveloped) (Text-figs 5 & 6).

The most constantly present pair of scutellar marginal setae throughout the

Tachinidae, unrepresented in a very few forms (e.g. some Minthoini).
basicosta. The small sclerite anteriorly at the wing base between the tegula
and the base of the costa (subepaulet).

The colour (whether clear yellow or orange instead of blackish brown) of this
sclerite is of minor taxonomic value in different parts of the family.

TACHINIDAE OF AUSTRALIA Il

bend of vein M. The forward curvature or angulation of the median vein (1)
where the bifurcation of WM, and M, occurs, or if there is no bifurcation then
the part of M where a forward change in its general direction occurs before it
attains the wing margin (cuwbitulus) (Text-figs 10 & 11).

The nature of the bend can be of great taxonomic importance (e.g. in
differentiating the Blondeliini with an open gently curving bend from the
Exoristini with a sharply angulate bend). A few Tachinidae lack the apical
part of the vein (i.e. M,) and there is therefore no bend, and some very rare
aberrant forms have only a very slight forward inclination of M.

cell R;. The wing cell distal to cross-vein v-m and enclosed by vein R,. , anteriorly
and veins M and M, posteriorly (apical cell) (Text-figs 10 & 11).

An important taxonomic character is provided by this cell and its associated
venation according to whether it is ‘open’ or ‘closed’. If the veins R4,,; and
M, reach the wing margin separately from each other then cell R; reaches to
the wing edge (Text-fig. 10) and is open; but when veins R,,, and M, coalesce
and form a short common stalk (‘petiole’) before reaching the wing margin the
cell is isolated from the edge of the wing and is closed (Text-fig. 11). In the
latter case the cell is described as ‘petiolate’ (prefixed ‘long-’ or ‘short-’ if
desirable). Intermediate forms occur in which the cell is closed just at the wing
margin.

The cell is open in the vast majority of Goniinae, Tachininae and Proseninae,
but is commonly closed and long-petiolate in Phasiinae (e.g. <Alophora,
Cylindromyia, Leucostoma, Euthera).

cerci. The inner median pair of articulated processes attached to the epandrium
of the male hypopygium (mesolobes, inner forceps, anal forceps) (Text-fig. 21).

Conformation and degree of fusion of the cerci have some taxonomic importance,
but relatively little at supraspecific level.

costa. The strong vein forming the fore margin of the wing.

costal sector. A segment of the costal length between any two successive veins
that end at the costa.

costal spine. A short strong seta on the costa at the apex of vein Sc, inserted
immediately before the costal ‘break’.

Usually undeveloped or very inconspicuous, if strong then normally not
exceeding v-m in length. Taxonomic value mainly at specific level.

cross-vein m-cu. The most distal wing vein connecting veins (7 and Cu, (posterior
cross-vein, hind cross-vein, m-m, M,) (Text-figs 10 & II).

The position at. which m-cu joins M (whether mid-way between 7-m and the
bend or nearer to the latter than the former) and the obliqueness of the vein
have some taxonomic importance (e.g. in Voriini the cross-vein is exceptionally
oblique in relation to the long veins and the wing in this tribe has a distinctive
appearance on this account).

cross-vein r-m. The very short length of vein connecting veins R,,, and M/, and
closing cell R, at its base (anterior cross-vein, R6) (Text-fig. 10).

In itself of no taxonomic use but a reference point for determining length
proportions on the long veins.

12 RK. W. CROSSKEY

discal setae. Setae standing medially or submedially on a specified surface (e.g.
scutellum or abdominal tergites).

Most often used with reference to strong erect setae standing centrally on
one or more of the abdominal tergites (the 1/D of Townsend).

distiphallus. The apical part of the aedeagus of the male genitalia (phallus,
aedeagus, preputwum) (Text-figs 22 & 23).

Of considerable taxonomic importance at various levels because of its diversity
of form. Sometimes bizarrely developed in whip-like or coiled form.

divaricate. Directed outwards from one another.

Used in particular of the ocellar setae when these curve neither forwards
(proclinate) nor backwards (reclinate) but curve outwards from each other
towards the eyes (e.g. in Paragonia).

dorsocentral setae. The two longitudinal rows of setae on the mesonotum outside
of the acrostichal setae (presutural + postsutural setae of Townsend) (Text-fig. 4).

Typically these are the strongest and most stable of the mesonotal setae
(excluding the presutural seta and the first supra-alar), but are weak or variable
in a few groups (notably Phasiini and Rutiliini). The number of dorsocentral
setae forming the presutural and postsutural complement is of great taxonomic
importance, particularly in the Goniinae. Many genera and tribes are completely
constant in their dc complement (e.g. all Sturmiini have 3 + 4 dc setae).

epandrium. The large curved plate of the male hypopygium that bears the cerci
and surstyli and is morphologically the ninth tergite (TG/X) (Text-fig. 23).

Apparently well developed in all Tachinidae but of limited taxonomic use

except for slight modifications in its vestiture.
epistomal axis. An imagined horizontal line through the head profile at the level
of the epistomal margin (oval margin axis) (Text-fig. 14). i
Head length at the epistomal axis in relation to length elsewhere (e.g. at the
antennal axis) can be significant in describing head form.
epistomal margin. The anteroventral edge of the epistome (oval margin).
epistome. The lower anterior part of the head below the face and between the
vibrissae (epistoma) (Text-figs 1, 2 & 12).

The development of the epistome is taxonomically important. Often there
is no epistome clearly differentiated from the face (e.g. Minthoini, Nemoraeini),
but in many forms there is a strongly developed epistome which curves forwards
from face and is easily visible in front of the vibrissal insertions when viewed
in profile (e.g. in many Alophora, most Rutiliini, Chaetophthalmus). In the —
Tachininae the presence of a prominently projecting epistome (visible in profile)
or a flat epistome (invisible in profile) provides a very important key character
separating groups of tribes. When the epistome is prominent its margin is
usually well below the level of the vibrissal insertions, and when exceptionally
pointed and projecting is usually referred to as subnasute.

erect. Standing upright in relation to the surface (said of hair or setae to contrast
their orientation with others that lie down) (cf. recumbent).
excavate. With a depression (said of abdominal Tr + 2).
The extent of the median dorsal depression of abdominal Tr + 2 is of great

TACHINIDAE OF AUSTRALIA 13

taxonomic importance at several levels in different parts of the family. Often
the depression extends backwards to reach the end of the segment, in which
case T1 + 2 is described as ‘excavate to its hind margin’, but in many forms
the depression clearly falls short of the hind margin or there is virtually no
depression at all (e.g. many Phasiinae, Doleschallini). When Tr + 2 is excavate
to its hind margin this condition often holds true of all the members of a tribe
(e.g. all Rutiliini, all Sturmiini); in other tribes (e.g. Blondeliini) the tergite
may be excavate to its hind margin in some genera but not others. In many
Prosenini the excavation is rather deep and narrow and the sides of Tr + 2
are rather strongly humped.

face. The anteromedian surface of the head below the antennae and between the
facial ridges (clypeus of Townsend) (Text-fig. 2).

facial carina. A strong median vertical ridge on the face separating the antennae
(facial keel) (Text-fig. 12).

The presence or absence of a facial carina, and its form when present, provide
important key characters for recognition of some forms. In most Tachinidae
the face is flat or only weakly raised medially, but a large facial carina is present
in many Proseninae (Prosena, Senostoma, Billaea and all Rutiliini). When a
carina is present it is typically flattened or convex on its anterior surface, or
prominently bulbous on its upper part and contracting to a sharper ridge
towards the lower end, but a sharp roof-like carina occurs in Euthera. <A true
facial carina is absent in Tachininae and Goniinae, but a weak swelling may be
present below and between the antennal bases (e.g. in Macrochloria).

facial profile. The anterior outline of the head between the antennal base and
the epistome when viewed in profile.

facial region. The face, facial ridges and parafacials collectively.

facial ridges. The flattened or raised strips differentiated on each side of the
face that separate it from the parafacials (facialia) (Text-fig. 2).

These are of great importance in taxonomy because of their variously modified
form (whether ridge-like lateral margins to the sunken face or flattened and
in the plane of the face), the extent of their visibility when the head is viewed
in profile, and the extent to which they bear hair or setae. In many aberrant
forms (e.g. Therobia, Myiotrixa) they are widened and flattened ventrally and
extensively haired, but typically they are at least slightly ridge-like and have
at least a few setulae immediately above the vibrissae (which in normal forms
demarcate the ventral ends of the ridges); in many genera of Tachininae and
Goniinae the ridges bear strong downcurved setae or setulae on most of their
height.

fascia. A transverse band on the dorsum of thorax or abdomen.

frons. The anterodorsal region of the head bounded by the eyes laterally and
extending from the ocellar triangle to the lunula; the interfrontal area and the
parafrontals collectively.

Typically the frons is well developed in both sexes, though usually narrower
in the male than the female, but is strongly reduced in some forms in which the
head is holoptic or almost so (e.g. in males of many Formosia, Ormiini and

14 R. W. CROSSKEY

Palpostomatini or in both sexes of many Phasiini). The frons is of equal width
in both sexes in all Siphonini and many Phasiinae, but rarely in other groups.
frontal setae. The setae of the paired rows of setae standing on the inner edges

of the parafrontals (Stivnborsten of Mesnil) (Text-fig. 3).

These setae are strongly or moderately developed in most Tachinidae but
may be weak and hair-like in Rutiliini and Phasiini and occasionally in other
groups. The rows may be partially obliterated in forms with holoptic or
semi-holoptic heads. Normally the setae of each row are directed inwards so
they cross medially (cruciate frontals), but in a few forms some or all of the
frontal setae are directed upwards and backwards (reclinate frontals). In
Phasiinae and Proseninae the rows of frontal setae extend downwards only to
the level of the antennal base or but slightly beyond, but in most Tachininae
and Goniinae the rows reach downwards at least to the level of the middle of
the second antennal segment and often beyond (in the latter case the rows
extend well down on the parafacials where, particularly in males, they may be
irregularly doubled, e.g. in Voriella).

gena. The lateroventral part of the head between the bottom of the eye and
the peristome or oral cavity (cheek, jowl, peristom of Mesnil) (Text-fig. 1).

The depth of the gena from the lowest point of the eye to the ventral margin
of the head when seen in profile can provide an important character, especially
its proportion in relation to eye height or width of the profrons (e.g. much narrower
in Carceliini than in other tribes of Goniinae).

genal dilation. The swollen haired part of the gena continuous with the postbuccae
and occiput (Occipitale Erweiterung auf dem Peristom of Mesnil) (Text-fig. 2).

Commonly the genal dilation occupies most of the gena and reaches well
forwards towards the vibrissal angles, but in some forms is reduced or virtually
absent (in which case the gena is bare or mainly so). The genal dilation is
normally extensively haired and sometimes bears a few stronger setae.

hair fascicles. Dense, often convergent, tufts of hair on the abdominal venter.

These are a secondary sexual character of males and occur mainly in a few
genera of Goniinae. When present they are normally restricted to the venter
or sides and venter of the fourth tergite (e.g. Palexorista) or of the fifth tergite
(e.g. Paropsivora, Zosteromeigenia), but occasionally are present on two successive
tergites.

humeral calli. The paired convexities forming the anterolateral corners of the
thoracic dorsum (humeri, posterior pronota, shoulders) (Text-fig. 4).
humeral setae. The setae standing on the humeral calli (Text-fig. 4).

The number and arrangement of these setae, particularly whether the main
ones stand in a straight line across the callus or in a triangle, is taxonomically
important at several levels. Usually each callus bears at least two differentiated
humeral setae, but there may be only one or even no clearly differentiated seta
(e.g. in Phasiini).

infrasquamal hairs. A group of minute hairs or setulae on the mediotergite
adjacent to the base of the lower calypter (Text-fig. 7).
Presence or absence of these hairs is sometimes a useful minor taxonomic

TACHINIDAE OF AUSTRALIA 15

character (but great care is often needed in examining them as they are easily
overlooked). They occur haphazardly in various genera in all the subfamilies
but are of rare occurrence in Goniinae (mainly confined to some Blondeliini)
and Proseninae (but occurring at least in some Chetogaster).

inner vertical setae. The main pair of erect setae standing on the summit of the
vertex (Innere Stirnborsten of Mesnil) (Text-fig. 3).

This pair of setae is almost universally present in Tachinidae, though it is
sometimes weak and hair-like in Phasiinae and some aberrant tribes. The two
setae are most often subparallel to each other but are sometimes directed inwards
and crossed, especially in some Tachininae. Subparallel inner verticals are the
rule in Goniinae though genera with cruciate inner verticals occur (e.g.
Phorocerosoma).

interfrontal area. The median area of the frons between the parafrontals
(interfrontalia, frontalia, frontal vitta, Stirnstreifen of Mesnil) (Text-fig. 2).

In forms in which the head is holoptic or the eyes strongly approximated
the interfrontal area is extensively obliterated and the parafrontals meet in
part in the mid line of the frons.

intermediate abdominal tergites. The middle two of the apparent four main
abdominal segments when seen from above, i.e. tergites 3 and 4 collectively.

intra-alar setae. The setae of the mesonotum standing external to the dorso-
central setae and approximately in line with the middle of the postalar
callus (Text-fig. 4).

The number and arrangement of these setae are of great taxonomic
importance. The most complete complement (excluding the occasional haphazard
supernumerary) is 1 + 3 ia setae and is found in the great majority of Goniinae.
The presutural intra-alar is most often absent in the other subfamilies but is
present, for example, in Voriini and Campylochetini (which have a I + 3 2a
complement like most Goniinae) and sometimes elsewhere. Many Phasiinae
and a few Tachininae have two strong posterior intra-alars that are widely
spaced with the anterior one close to the transverse suture, e.g. Leucostoma,
Cylindromyia, Zita, Leverella; this arrangement of two post ia is fundamentally
different from that found in forms normally possessing three but occasionally
lacking the anterior one (in which case the two remaining post ia stand near
each other and the front one is remote from the transverse suture). A few
forms completely lack intra-alar setae (e.g. some Phasiini and Ormiini) or they
may be missing in isolated specimens of species normally possessing at least
one. In Cylindromyia the anterior one of the two post ia is enormously strong
and stands out of alignment with the hind one and unusually close to the
pre-alar seta.

last section of Cu,. The part of vein Cu, between its junction with m-cu and its
apex at the wing margin.
lateral scutellar setae. One or more pairs of marginal scutellar setae standing
between the basal and subapical or apical pairs (Text-fig. 5).
These setae are frequently absent, and when present are usually weaker than
the basals and subapicals.

16 R. W. CROSSKEY

lower calypter. The larger ventral one of the pair of membranous lobes at the
wing base rigidly attached at its base to the suprasquamal ridge (squama,
thoracic squama, Thorakalschiippchen of Mesnil).

Taxonomic characters of the lower calypter lie in its shape and vestiture,
and at specific level in colour. It is well developed in all Tachinidae, and in
some is grossly enlarged (e.g. in an undescribed Leucostomatine from
Madagascar the lower calyptrae of the male are so enormous that they completely
hide the abdomen). In some forms (e.g. Palpostomatini) the lower calypter is
subcircular posteriorly and diverges from the scutellum but in many it is very
broad posteriorly and its inner edge abuts close to the scutellum (e.g. Sturmiini
and many other Goniinae, many Phasiini). In a few Goniinae (Ethillini, some
Winthemiini, some Carcelia) the outer edge of the lower calypter is bent abruptly
downwards. Normally the upper surface is bare but fine erect hairing is present
in a few forms and provides a useful key character (e.g. Nemoraea, some Senostoma,
some Rutilia (Chrysorutilia)).

lunula. The crescentic sclerite between the antennal bases and the frons (frontal
lunule) (Text-fig. 2).

Useful as a descriptive landmark, but otherwise of little value. Bare in all

Tachinidae.
marginal scutellar setae. The strong setae around the edge of the scutellum
collectively.

The total number of pairs of these setae can provide a useful key character.
In a few forms the marginal setae are more numerous than usual, slightly or
strongly spiniform, and not clearly differentiated into basals, laterals and
subapicals (e.g. Formosia, Rutilia).

median marginal setae. Setae standing on the middle part of the hind margin
of an abdominal tergite (the MM of Townsend).

mediotergite. The posterodorsal declivity of the thorax below the postscutellum
and between the halteres (Jateral postscutellar plates + basiscutellum of Townsend)
(Text-fig. 7).

Not used in taxonomy except for presence or absence of fine hairs on the
mediotergite near the base of the lower calypter (infrasquamal hairs, q.v.).
Shape might have taxonomic value but has been insufficiently investigated.
In some forms the middle part immediately ventral to the postscutellum (i.e.
the basiscutellum of Townsend) is differentiated by a distinct line on each side
from the lateral parts (i.e. the lateral postscutellar plates of Townsend), but in
others (e.g. Doleschalla) there is little sign of such differentiation.

mentum. The sclerotized shaft or main plate of the proboscis (figs 12 & 13 in
Crosskey, 1973).

Length and shape of the mentum of the proboscis have some taxonomic use,
e.g. in Rutiliini in which the mentum in some forms seen in profile is distinctly
tapering while in others it is parallel-sided.

mesonotum. The prescutum and scutum together (Text-fig. 4).

Markings of the mesonotum when present, e.g. vittae or spot-patterns, have

some taxonomic usefulness. Strictly, the mesonotum is formed of a scutum

TACHINIDAE OF AUSTRALIA 17

subdivided by the transverse suture, but the terms prescutum and scutum
remain useful in taxonomy.

mesopleuron. The upper anterior part of the wall of the thorax between the
anterior spiracle and the wing base (anepisternite) (Text-fig. 7).

Of little use taxonomically. Presence or absence of a dense pollinose spot
on the centre of the mesopleuron is occasionally useful at specific level (e.g.
in Rutiliini). A small anterodorsal mesopleural seta may be developed but is
little used taxonomically.

notopleuron. A small subtriangular area differentiated laterodorsally on the
thorax between the humeral callus and the wing base and external to the
prescutum (Text-figs 4 & 7).

Of very little use in taxonomy except as a minor aid in distinguishing between
a few Proseninae. In Doleschallini the notopleura are not noticeably
differentiated from the prescutum, and in a few Rutiliini (e.g. subgenus
Neorutilia) the posterior part of the notopleuron is unusually swollen and
knob-like.

notopleural setae. The setae standing on the notopleuron (Text-fig. 4).

In very nearly all Tachinidae these number two, one anterior and one posterior.
Rarely additional setae are developed, e.g. in the Philippines species of Rutilia
(Chrysorutilia) in which the posterior knob-like swelling of the notopleuron
bears two strong setae (in place of the usual single posterior notopleural).

occiput. The posterior surface of the head excluding the postorbits (Hinterkopf
of Mesnil) (Text-fig. 1).

occipital setulae. Recumbent black setulae or strong hairs on the upper occiput
behind the postocular row (schwarzen Mikrochdten hinter den Postokularbértschen
of Mesnil).

Presence or absence of these setulae is often a useful taxonomic character at
various levels.

ocellar setae. The pair of setae standing very near the ocelli on the ocellar triangle
or tubercle (Text-fig. 3).

Presence or absence, orientation, and occasionally the precise position of
the insertions relative to the ocelli are of taxonomic value and provide useful
key characters. When present the setae are most often curved forwards and
downwards (proclinate ocellars), but occasionally backwards (reclinate ocellars)
(e.g. in Gonia, Elpe, Leucostoma); in a few forms they curve directly outwards
towards the eyes (divaricate ocellars) (e.g. in Paragonia).

ocellar triangle. The subtriangular area of the vertex on which the ocelli are
placed (ocellar plate, vertical triangle, ocellarium) (Text-fig. 2).

| Typically the ocellar triangle is slightly raised in relation to the rest of the

_ vertex, and in forms with holoptic or subholoptic heads the ocelli are on a very

|

|

prominent ocellar tubercle (e.g. most male Palpostoma). Ocelli are almost
universally present in Tachinidae, but are vestigial or absent in some Ormiini
(e.g. Therobia).

_ outer vertical setae. A pair of outwardly curved setae standing laterally on the

vertex close to the eyes (Text-fig. 3).

B

18 R. W. CROSSKEY

These setae are weak or absent in many forms, and when present are usually
only strong in females. They are present in females of Goniinae, but usually
absent or represented by mere hairs in males (in the Siphonini both sexes have
strong outer verticals).

palpi. The paired forwardly directed appendages of the proboscis arising between
the base of the mentum and the oral cavity.

Presence or absence of palpi, and shape and size when present, often provide
useful key characters. Palpi are usually present (virtually always in Goniinae)
but may be totally absent as in Cylindromyia or vestigial (e.g. Linnaemya,
Chaetophthalmus, some Stomatomyia). Exceptional development sometimes
occurs in which the palpi project far beyond the epistomal margin (e.g.
Exechopalpus) or are greatly swollen and clubbed (e.g. females of Rutzlotrixa).

parafacials. The paired anteroventral areas of the head separating the eyes from
the facial ridges and lying between the parafrontals and the genae (parafacialia,
Wangen of Mesnil) (Text-figs 1 & 2).

These areas are taxonomically important for their vestiture and breadth in
relation to, for instance, the width of the face or the antennae. Most often
the parafacials are bare, but they may be haired to a varying extent or may
carry strong bristles. In many forms the descending rows of frontal setae
reach down on to the upper parts of the parafacials, but these setae are discounted
when the parafacials are described as bare (in descriptive practice the ‘parafacials’
normally means those parts of them lying ventrally to the lowermost frontal
setae).

parafrontals. The paired areas forming the outer parts of the frons and abutting
against the eyes (orbits, fronto-orbital areas, parafrontalia) (Text-figs I & 2).

Normally the parafrontals are separated from each other by the interfrontal
area, but in some forms they meet each other in the mid-line of the frons and
partially or wholly obliterate the interfrontal area. Usually when they meet
the head is holoptic or nearly so, but a very few forms with very broad frons
have parafrontals that not only meet in the mid line but have lost most or all
of the line of union so that the entire frons is formed of completely coalesced
parafrontals (e.g. males of most Heterometopia). Hairing of the parafrontals
can be of minor taxonomic usefulness at specific level.

peristome. The lower margin of the head around the buccal opening.

The term is used by Mesnil for the whole of the area here termed the gena.

peristomal setae. Lower marginal setae of the head standing on the peristome
(Text-fig. 3).

petiolate. Provided with a petiole (q.v.) (said of wing cell R;).

petiole. The short length of wing vein formed by the union of the apices of veins
R,,, and M, in wings in which cell R, is closed before the wing margin. See
under cell R; above.

pleurotergite. The convex area of the side of the thorax lying above the hind
spiracle and between the pteropleuron and the mediotergite (sguamopleuron,
supra-spiracular convexity).

This area is bare (i.e. has only a nap of microscopic pubescence) in nearly all

|
|

TACHINIDAE OF AUSTRALIA 19

Tachinidae but has taxonomic importance as a taxonomic feature in a few
forms in which it bears long dense hair (e.g. Microtropesa and Paratropeza).

pollinose. With a covering of pollinosity (q.v.).
pollinosity. A nap-like covering to any part of the body formed by closely

aggregated ultramicroscopic pubescence.

The extent, density and colour of pollinosity can provide taxonomic characters
and are largely responsible for the general appearance of many Tachinidae.
Banded or spotted patterns are typically produced either by islands of dense
pollinosity surrounded by less pollinose areas, or alternatively by non-pollinose
islands in areas that are mainly thickly coated with pollinosity (e.g. the
conspicuous spot patterns of Amphibolia). When pollinosity is not evident
at normal magnifications of the entomological microscope (up to X I00 or x 150)
or is very inconspicuous on any area then the area is said to be ‘non-pollinose’.
When the cuticle is dark, non-pollinose areas are typically metallic and the extent
of metallic colour can be important taxonomically (e.g. in Rutiliini).

postabdomen. The part of the abdomen including the terminalia lying beyond

segment 5 (the last segment normally readily visible).

In many forms much of the postabdomen is concealed and tergite 5 forms
the end of the immediately visible abdomen. In some forms the postabdomen
is more conspicuously developed than normal and may be distinctly recurved
under the main part of the abdomen (preabdomen) (e.g. Cylindromyiini) and
may bear elaborate modifications associated with oviposition (e.g. various
forcipate processes in female Phasiinae).

postalar calli. The paired convexities forming the posterolateral corners of the

thoracic dorsum (Text-figs 4 & 7).

postalar setae. The strong horizontal setae standing on the postalar calli

(Text-fig. 4).

In nearly all Tachinidae there are two strong postalar setae, but in all
Rutiliini there are three or more (sometimes as many as six or seven) strong
postalars, the distinction between three, and four or more, being very important
in taxonomy of the group. Outside of Rutiliini the development of a strong
third postalar seta is rare (but occurs, e.g., in some Microtropesa). Rarely only
a single strong postalar seta is developed (e.g. in some Phasiini).

postalar wall. The vertical declivity forming the outer surface of the postalar

callus (Text-fig. 7).

This area is of taxonomic value in the Rutiltini where the postalar wall bears
a dense tuft of hair in Formosia and Formodexia Crosskey that is absent in
other genera. These genera appear to be the only forms represented in the
Tachinidae with such hair (in all others the postalar wall is bare or at most has
only one or two hairs on its extreme upper part immediately below the rounded
haired outer lip of the callus).

_postbucca. A vaguely defined area between the gena and the occiput forming

|

the posteroventral corner of the head (posigena) (Text-fig. 1).

posteroventral declivity of the thorax. That part of the thorax laying between

the base of the abdomen and the insertions of the hind coxae (Text-fig. 7).

20 R. W. CROSSKEY

In most Tachinidae the hind coxae are rather close to the abdominal base
and the posteroventral declivity of the thorax consists of a median membranous
area flanked by curved sclerotized areas (the metapleura). In a few forms
(e.g. Cylindromyia, Doleschalla) the abdomen and the metacoxae are much
more remote from each other than is usual and the entire posteroventral
declivity of the thorax forms a fully sclerotized bridge between them; this forms
an important taxonomic and key character.

posthumeral setae. One or more setae standing anterolaterally on the prescutum
immediately behind the humeral callus (Text-fig. 4).

Development of these setae is rather variable and their taxonomic usefulness
very limited. The pf setae are usually most strongly developed in Goniinae,
in which two moderately strong posthumerals are normally present on each
side, and least strongly represented in Phasiinae and Proseninae in which they
may be absent or hair-like.

postocular row. The regular row of setulae or long hairs on the upper and outer
edges of the occiput immediately behind the eyes (Text-fig. 3).

postorbits. The narrow laterodorsal strips of the head lying between the eyes
and the postocular rows (Text-fig. 1).

postscutellum. The transverse swollen lobe below the scutellum and dorsal to
the mediotergite (subscutellum, infrascutellum, postnotum) (Text-fig. 7).

This is almost universally present in Tachinidae and strongly convex. It
is very weakly developed in a few forms (e.g. Palpostomatini) and is strongly
produced backwards, squared and flattened in some Phasiini (e.g. Campbellia,
Alophora subg. Mormonomyia). The postscutellum is always bare.

postsutural. Behind the transverse suture of the mesonotum.

pre-alar seta. A seta standing at the extreme anterolateral corner of the scutum
anterior to, and approximately in line with, the supra-alar setae (first swpra-alar
seta of authors) (Text-fig. 4).

This seta is of the very greatest importance in tachinid taxonomy because
of its variability in size, and, to a lesser extent, because of its presence or absence.
The seta is universally present in the Goniinae (though it may be very weak)
and may be either small (shorter than the first post ia seta or at most subequal
to it) or very large (longer and stronger than the first post ia seta and usually
also longer than the first post dc seta), and this distinction holds true almost
without exception for the differentiation of two groups of Goniine tribes. In
Phasiinae, Proseninae and most Tachininae the pre-alar seta is of the small
type, and is sometimes absent (e.g. in Phasiini, some Minthoini and sometimes
haphazardly in Proseninae).

preapical (discal) scutellar setae. A pair or more of recumbent or semi-
recumbent setae standing on the scutellum in advance of the marginal setae
(Text-fig. 5).
prescutum. That part of the mesonotum lying anterior to the transverse suture
(Text-figs 4 & 7).
Morphologically this is the anterior part of a divided scutum but the term

TACHINIDAE OF AUSTRALIA 21

remains useful in taxonomy. The prescutum as such has little use in taxonomy
except for its patterning and pollinosity.

presutural. In front of the transverse suture of the mesonotum.

presutural seta. A single seta standing on the outermost part of the prescutum
near the notopleuron and approximately in line with the pre-alar and supra-alar
setae (presupra-alar seta of Townsend) (Text-fig. 4).

This is the most consistently stable and universally present thoracic seta in
the Tachinidae, and on this account is not used in taxonomy. It is very nearly
always present at least as a differentiated hair, and in the overwhelming majority
of forms is an extremely strong seta even when other parts of the chaetotaxy
are much reduced.

prevertical setae. A pair of outwardly directed setae standing on the upper ends
of the parafrontals.

These setae occur mainly in females of certain Tachininae and are very rarely
present in Goniinae (though found in some Spfoggosia and Baumhaueriina, at
least). They appear to represent a pair of orbital setae that curve outwards
over the margins of the eyes instead of backwards towards the vertex (as is
typical in orbital setae).

proclinate. Directed forwards, or forwards and downwards.

proclinate orbital setae. A pair or more of strong downwardly directed setae
standing externally on the parafrontals near the eyes (fronto-orbital setae, Aussere
Orbitalborsten of Mesnil) (Text-fig. 3).

These setae may be absent in both sexes, but typically two pairs are present
in females and none in males. In some forms (e.g. all Siphonini) they are present
in males as well as females. Occasionally a series of several (up to six or seven,
rarely more) pairs may be present in both sexes (e.g. Halydaia, Ceracia).

profrons. A weakly defined area where the parafrontals and the parafacials
meet (Text-figs 1 & 13).

Typically the area of the profrons is somewhat projecting and seen in profile
is the widest part of the head that is visible anterior to the eye. The width
of the profrons seen in profile in relation to the depth of the gena is occasionally
a useful character.

propleural seta. A seta (sometimes duplicated) standing on the extreme
anteroventral corner of the thorax (Prdstigmdtikalborste of Mesnil) (Text-
fig. 7).
This seta, even if only hair-like, is differentiated in almost all Tachinidae
and has very limited taxonomic use. It is totally absent in a few Goniinae
: (e.g. Anacamptomyia, Euvespivora, some Carcelia s.l. species), rarely elsewhere.
propleuron. The flattened area of the front edge of the thorax below the humeral
callus (Text-fig. 7).

This area has considerable taxonomic importance and provides an easily
|) seen key character according to whether it is bare (as is most often the case)
or haired. The area never has strong setae. Hairing of the propleuron occurs
sporadically throughout the Tachinidae, but particularly is present in all
Rutiliini and many Tachininae (e.g. Campylochetini). In the Phasiinae the

22 Ri Wi CROSSIGE ¥

propleuron is almost always bare, and in the Goniinae hairing of the propleuron
is almost entirely confined to certain Blondeliini and Neaerini (the propleuron
appears to be consistently bare in the tribes of Goniinae in which the pre-alar
seta is very large).

prosternal membrane. The extensive membranous area on either side of the
prosternum (Text-fig. 9).

This membrane is only of taxonomic value according to whether it is bare
(as in the overwhelming majority of Tachinidae) or haired. Hairs on the
prosternal membrane occur mainly in Rutilia subg. Chrysorutilia and a few
allied Rutiliini, but elsewhere are of very rare occurrence (found, for example,
in some Chaetophthalmus species).

prosternum. The strong ventral plate of the thorax lying between and in front
of the fore coxae (Text-fig. 9).

One of the most important taxonomic characters in the Tachinidae is
provided by the presence or absence of vestiture on the prosternum. The
vestiture when present consists of fine hairs or a few strong setulae on each
side of the middle part of the prosternum which are directed outwards or
downwards, or of some soft hair on the anterior corners (as in some Rutilia
and some Chaetophthalmus species). The prosternum is totally bare in
(apparently) all Phasiinae, almost all Proseninae and the great majority of
Tachininae, but is almost universally haired or setulose in the Goniinae (even
if the vestiture consists only of a single hair on each side); a few Goniinae have
the prosternum bare, these including several genera of Blondeliini and
Blepharella in the Sturmiini.

prostigmatic seta(e). One or more strong setae anterolaterally on the thorax
below the anterior spiracle (mesopleurospivacular setae of Townsend, Substig-
matikalborsten of Mesnil) (Text-figs 7 & 8).

One or more of these setae almost always present and directed upwards.
The genus Peribaea (tribe Siphonini) is very exceptional in having two equally
strong prostigmatic setae of which the upper one is directed upwards and the
lower one strongly downwards (an arrangement apparently found nowhere else
in the family) (Text-fig. 8).

pteropleural seta. A seta standing on the upper edge of the pteropleuron
(Text-fig. 7).

Presence or absence of this seta can provide an important character at various
levels. Sometimes it is only weakly differentiated from the general pteropleural
hairing and sometimes is duplicated. The seta is absent or very weak in nearly
all Phasiinae and Proseninae and strong in nearly all Goniinae; in the Tachininae
it is developed to various degrees in different genera or tribes, and if strong its
actual size in relation to other structures is sometimes taxonomically useful
(e.g. in African Linnaemya).

pteropleuron. The area of the side of the thorax immediately behind the
mesopleuron and below the wing base (Text-fig. 7).

The only use of this area in taxonomy (apart from the pteropleural seta, q.v.)

is for the extent of its hair vestiture. Normally the anterior half of the

~

TACHINIDAE OF AUSTRALIA 23
pteropleuron (and sometimes much of the lower part) is bare but in a few taxa
(e.g. the subgen. Chrysorutilia) the hairing extends further forwards of the
pteropleuron than is usual.

reclinate. Directed backwards, or upwards and backwards.
reclinate orbital setae. One or more pairs of erect or backwardly directed setae
standing on the upper parts of the parafrontals (Text-fig. 3).

The number of pairs of these setae can provide an important taxonomic
feature, as in Sturmiini in which genera with one pair are rather sharply
differentiated from genera with two or more. The setae are best differentiated
in the Goniinae and some Tachininae, but may be absent in males of some
forms though present in females (e.g. Winthemia) or may be very imprecisely
differentiated from the rows of frontal setae. Reclinate orbital setae are absent
or at least indefinite in Phasiinae and Proseninae. In genera in which males
have a single pair of strong isolated reclinate orbitals the females may show
two pairs, but have the lower pair weaker than the upper (main) pair and
situated closer to each other.

recumbent. Lying down or appressed to the surface (said of hairs or setae to
contrast their orientation with others that stand upright) (cf. erect).

scutellum. The large hemispherical or subtriangular lobe of the thoracic dorsum
behind the scutum.

The shape and degree of flattening of the scutellum, and the nature of its
hairing, provide minor taxonomic characters. }

scutum. That part of the mesonotum lying posteriorly to the transverse suture
(Text-figs 4 & 7).

Morphologically this is only the posterior part of a divided scutum but the
term remains useful in taxonomy. The scutum as such has little use in taxonomy
except for its patterning and pollinosity.

second aristal segment. The more distal one of the two basal segments of the
arista.

This segment varies in its length (to a greater extent than the first aristal
segment) and can provide a useful feature.

second costal sector. That segment of the costa lying between the apices of veins
Se and R, (Text-fig. 10).

The presence or absence of small fine recumbent hairs along the ventral
surface of this costal sector provides a useful taxonomic and key character at
various levels in different parts of the family. The hairs when present are
additional to, and quite distinct from, the marginal costal setulae. Sometimes
whole tribes have one condition or the other (e.g. second costal sector bare below
in all Rutiliini or haired below in all Thelairini), but elsewhere the character
may be generic only (e.g. in Blondeliini).

seta, setula. Not differentiated from each other on any absolute criterion, but
seta generally reserved for any strong and obvious bristle forming a particular
component of the formalized chaetotaxy, and setula for a small stiffened hair.
Setulae are commonly serially arranged, as along the costal margin, down the
facial ridges or in the postocular row.

24 R. W. CROSSKEY

spiniform setae. Those setae that are exceptionally thickened and spine-like and
lack the even tapering from base to apex of typical setae.

These occur commonly in Rutiliini and Tachinini and to a lesser extent elsewhere
(e.g. some on the scutellum of Goniini).

sternite 5. The deeply divided last visible abdominal sternite of the male
(Text-fig. 20).
sternites. The sclerotized plates of the mid venter of the abdomen (Text-fig. 20).

The extent of exposure or concealment of the sternites by the ventral ends
of the tergites can provide a useful taxonomic character. The sternites may
be completely exposed in their surrounding membrane (as in many Phasiini)
or completely concealed by overlapping of the tergites (as in Goniinae), and
intermediate conditions occur in which the anterior sternites are mainly or
fully concealed and the fifth sternite and part of the fourth exposed. Vestiture
of the sternites is occasionally of minor taxonomic use.

sternopleural setae. Setae standing on the wpper part of the sternopleuron
(Text-fig. 7).

The number and arrangement of these provide very valuable characters.
Rarely they are totally absent or only one (posterior) one is present, but typically
there are two or three (arranged I + I or 2+ 1). Four or (very rarely) more
occur in some Goniinae and a few Tachininae and the arrangement of these
may be significant (e.g. a characteristic group of genera of Sturmiini has the
sternopleurals constantly arranged 2 + 2). When three are present the size of
the lowermost seta in relation to the others may be useful.

sternopleuron. The large lateroventral subtriangular area of the thorax between
the fore and mid coxae and below the mesopleuron and pteropleuron (Text-fig. 7).

Normally important only for its vestiture and this has very limited taxonomic
use. A characteristic row of fine regular recumbent hairs in front of the mid
coxa is present in Actia, and the ventralmost hairing may be modified into strong
spiniform setae.

subapical scutellar setae. The pair of setae inserted posterolaterally on the
scutellar margin (Text-figs 5 & 6).

These setae, together with the basals, are the most universally present setae
on the scutellum and typically are the strongest pair. Their distance apart
from each other at the base in relation to the distance between the insertion of
one subapical seta and the basal seta of the same side of the scutellum provides
an important character. Also important is the direction of the subapicals,
e.g. whether diverging from each other as in Blondeliini or converging and
crossing at their tips (enclosing the apical setae) as in Siphonini and Neaerini.

sublunular bulla. A knob-like swelling between the antennal bases immediately
below and continuous with the lunula.

This occurs mainly in Phasiini, where in Alophora and allied genera the bulla
is often polished and shining.

supra-alar area. The outer edge of the scutum above the wing base.
supra-alar setae. The outermost setae standing on the scutum (postsupraalar
bristles of Townsend), other than the pre-alar seta (Text-fig. 4).

TACHINIDAE OF AUSTRALIA 25

These are normally two, one behind the other, with the anterior one very
strong. The anterior one is, with the presutural seta, one of the most stable
and universally present setae in the tachinid chaetotaxy. Sometimes one or
two supernumerary supra-alars may be developed in addition to the normal
two, or the posterior supra-alar may be absent or hair-like. The second or
posterior sa seta is always present in Goniinae but is absent in such tribes as
the Phasiini and Palpostomatini and often in Ormiini.

Some authors consider the pre-alar seta (pra) as the first of the supra-alar
setae and number the others accordingly (e.g. the posterior supra-alar is then
the third supra-alar), but this is an undesirably confusing terminology. When
both the pre-alar and the posterior supra-alar are absent (as in Phasiini and
Palpostomatini) the anterior supra-alar is the only seta present on the supra-alar
area and therefore stands very conspicuous and isolated.

suprasquamal ridge. The weakly sclerotized area at the base of the postalar
wall to which the lower calypter is attached marginally (tympanic ridge of
Townsend) (Text-fig. 7).

This structure is taxonomically useful according to whether it is bare or
haired. The suprasquamal ridge is bare in all Phasiinae and Goniinae, and
in the Proseninae hairing is confined to the Rutiliine genera Rutilia, Amphibolia
and Chrysopasta; the ridge is apparently bare in all Tachininae except Servillia
jakovlewti (Portschinsky).

surstyli. The outer pair of articulated processes attached to the epandrium of
the male hypopygium (paralobes, outer forceps, tenth sternite lobes of Townsend)
(Text-figs 21 & 23).

The form of the surstyli is of great taxonomic value at specific level but they

have relatively little taxonomic use at higher levels.

tegula. The small anterior sclerite at the exteme base of the wing before the
basicosta (epaulet).

tergites. The main segmental plates of the abdomen forming almost the entire
circumference of each segment.

The tergites are taxonomically important in so far as their individual and
collective forms give shape to the abdomen and determine, for example, whether
it is elongate subcylindrical as in Cylindromyia and Doleschalla, or rather broad
and flattened as in many Phasiini and Rutiliini, or subovate as in the majority
of forms. In the Goniinae the ventral ends of the tergites meet in the mid-
line of the abdomen, but in many forms in the other subfamilies they do
not meet each other and the sternites are then exposed. Lengths of the
tergites relative to each other, special modifications such as the depression of
T5 found in many Rutiliini or the keel-like form of the abdominal venter in
some Blondeliini, and pollinosity and pattern on the tergites can all provide
useful characters.

third costal sector. That sector of the costa lying between the apices of veins
kK, and R, . 5.

_ transverse suture. An impressed line across the mesonotum dividing it into

anterior and posterior parts (prescutum and scutum, q.v.) (Text-figs 4 & 7).

26 KR. We CROSSIGEY

upper calypter. The smaller dorsal one of the pair of membranous lobes at the
wing base (alar squama).

This has little use in taxonomy but is of note in Prodiaphama (tribe Rutiliini)
in which, when the wings are closed, it is as long as or almost as long as the
lower calypter. In other Tachinidae it is normally much shorter.

venter. The lower surface as a whole (especially of the abdomen).
vertex. The upper surface of the head between the eyes and around the ocellar
triangle (Text-figs 1 & 2).

This area is precisely delimited laterally by the eyes but it is rather vaguely
defined from the frons and occiput. The width of the vertex varies greatly
according to how closely the eyes are approximated and is typically wider in
females than males. The width when seen from above in relation to total head
width provides important specific differences. When the head is fully holoptic
(e.g. In some Palpostomatini) all that remains of the vertex is a prominent
ocellar tubercle (see ocellar triangle).

vestiture. Setae and hairing collectively.
vibrissae. Paired setae inserted anteroventrally on the head at the lower ends
of the facial ridges and directed forwards (Text-fig. 3).

Typically the vibrissae are extremely strong and curve forwards and inwards
like a pair of tusks; often they are the strongest setae borne on the head. In
some forms they are reduced and scarcely distinguishable from the peristomal
setae (below them) or from the small setulae on the lower ends of the facial
ridges (above them). Short stubby non-crossing vibrissae occur in some forms.

vibrissal angles. The corners of the head, typically rather prominent, on which
the vibrissae stand (Text-fig. 1).

vibrissal axis. An imagined horizontal line through the head profile at the level
of the vibrissal insertions (Text-fig. 14).

vitta. A longitudinal band or line on the dorsum of the thorax or abdomen.

vittate. Marked with vittae.

wing venation. The rigid veins that support the wing membrane collectively.

This provides many important taxonomic characters at several levels,
according to the proportions shown by different vein lengths, the fusion or
non-fusion apically of R,,; and M,, the position of the bend of vein M in
relation to the wing margin, the evanescence of parts of veins (especially loss
of M,), the positions at which long veins abut on the costa, and to whether
certain veins reach or fail to reach the wing edge. In addition the extent and
presence or absence of setulae on certain veins can provide useful characters.
Setulae may occur on veins R,, Ry. 5, or Cu, but not on the other long veins
and never on the cross-veins; they are usually more extensive when present
on the upper surface of the wing but may occur ventrally also on R, and Ry+;
(especially on the node).

KEY TO AUSTRALIAN SUBFAMILIES OF TACHINIDAE

The satisfactory classification of the Tachinidae into subfamilies is a matter of
great difficulty and specialists are still far from agreed on the number and scope

:
|
|

{
|

TACHINIDAE OF AUSTRALIA 27

of the subfamilies that should be recognized and on the combinations of characters
that should be used to define them. Most taxonomists currently recognize relatively
few subfamilies (normally between four and six) but a large number of tribes, but
none of these family-group taxa are susceptible of simple and unequivocal definition
that will render their members immediately identifiable on straightforward key
characters. The existence of many aberrant forms, and of much convergent
evolution, compounds the classificatory difficulties still further and ensures that —
view them how we may — it is simply impossible to define the subfamilies succinctly
or to key them out (at least on the external morphology of the adults) with any
precision. Undoubtedly, however, recent and continuing studies on the male
genital structure, the female reproductive habit, the morphology of the larvae
(especially the first instar larvae), and of the host relations, are doing much to
enlighten specialists on the probable phyletic relationships, and hopefully this newly
acquired data may lead to more concrete subfamiliar definition in the course of time.
Meanwhile, it is possible to recognize many of the members of the more distinctive
subfamilies (Phasiinae, Goniinae) on their overall facies, even though it is extremely
difficult to fix their characters in an unexceptionable way that is satisfactory for
key construction, and a knowledge of the host relations of any tachinid parasite
can be of immense value in placing its subfamiliar identity -for example, any
Australian tachinid with a host in the Hemiptera belongs in the Phasiinae.

In the present work four subfamilies are recognized, the Phasiinae, Proseninae
(=Dexiinae), Tachininae and Goniinae, but it is freely admitted that the Tachininae
in the sense here used is a heterogeneous assemblage of forms in which two or three
distinct subfamilies ought probably to be acknowledged; but until a clearer picture
of the interrelationships of the included forms emerges it remains useful to treat all
the Australian tachinids that are clearly not either Phasiinae or Proseninae or
Goniinae as constituting one subfamily (for which the name Tachininae is nomen-
claturally correct).

With the difficulties in defining the subfamilies, and the confusing overlap in
many of their characters, it is impossible to construct a workable key even to the
small number of subfamilies here recognized that will permit each and every specimen
to be placed unequivocally in a subfamily. The key that follows is only a tentative
guide to the probable suprageneric grouping to which any specimen belongs. In
order to keep the key short and simplified (without long confusing couplets that
attempt to cover every exceptional or aberrant form) certain tribes have been run
out individually and sometimes there is more than one exit for a particular sub-
family. Some extremely poorly known forms of very uncertain subfamilial position
(e.g. Myiotrixa, Amphitropesa, Neximyia) have been omitted from consider-
ation, as to include them would so complicate the key as to negate much of its
value. When the names of taxa are juxtaposed this does not imply close phyletic
relationship.

I Ocelli absent. Prosternal region grossly inflated, visible from side view. [Probably
parasites of Orthoptera] . : : : é . Ormiini (Tachininae, part) (p. 57)
— Ocelli present. Prosternal region normal, not visible in profile. : ‘ : 2

28

on

K. W. CROSSKE Y¥

Face with a heavy facial carina extending from epistome to lunula between the
antennae and convex, bulbous or flattened on much or all of its anterior surface
(Text-fig. 12). Eyes bare. [Parasites of Coleoptera] . PROSENINAE (part) (p. 41)

Face without such a facial carina, usually flat or sunken, if weakly carinate just
below antennal bases (Macrochloria) then eyes haired, if bullate between antennal
bases only (Alophora) then cell R; long-petiolate (but sharp ridge-like facial carina
on whole face present in Eutherini) . : : : 3

One supra-alar seta (Text-fig. 54). Pre-alar se Bee (eae Saale. Usually
one post ia seta or none. Head often holoptic or almost so and upper eye facets
often conspicuously enlarged. Eyes always bare : 4

Two or more supra-alar setae (Text-figs 56-63) (hindmost sss wea sercae
may be absent in Minthoini). Pre-alar seta almost always present (sometimes
hairlike). Normally two or three post ia setae. Head never fully holoptic and
eye facets never greatly enlarged. Eyes bare or haired . : : : : 5

Lower calypter very broad, straight or concave on its hind margin. Prosternum
bare. Labellae without palpiform processes. Not more than one fost ia seta.
[Parasites of Hemiptera] . . PHASIINAE (part) (p. 29)

Lower calypter subcircular posteriorly and ‘strongly diverging from the scutellum.
Prosternum on each side with a long seta directed downwards. Labellae usually
with a distinct pair of palpiform processes. Normally two post ia setae. ([Para-
sites of Coleoptera) . ; : Palpostomatini (Tachininae, part) (p. 53)

Two post ia setae present standing widely apart from each other with the anterior
one not far from the transverse suture, the setae either subequal in size or the
antevioy one enormous and much stronger than the posterior one (Text-figs 56 & 58).
Either the hind coxae very widely separated from the abdominal base by a deep
completely sclerotized bridge, or apex of 2 abdomen with forceps-like processes,
or both. Postabdomen usually strongly recurved under the preabdomen. Wing
with cell R, closed and long-petiolate (Text-figs 76 & 78). [Parasites of Hemiptera]

PHASIINAE (part) (p. 29)

Two or more (usually three) post ia setae present which increase in size towards the
hindmost (Text-figs 57,59-63), if only two are present then the anterior one far
from the transverse suture (except in Parerigonini and in Minthoxia in which two
widely spaced subequally strong post ia setae present, but then other characters
not fitting). Hind coxae and abdominal base not very remote from each other,
area between them membranous medially; end of 2 abdomen never forcipate.
Postabdomen not distinctly recurved under preabdomen. Wing with cell R;
usually open, a few exceptions (e.g. Euthera). [Not parasites of Hemiptera, except
probably Eutherini| : F . 6

Wings with a pattern of eee eee black panes (est fig. a5) Celle
with long petiole (Text-fig. 77). Face with a very prominent sharp median ridge
up its whole height (Text fe 24). Antennae exceptionally long, their apices
reaching beyond the epistomal margin. [Probably parasites of Hemiptera, no
Australian records | F Eutherini (Phasiinae, part) (p. 39)

Wings without any bold nae pattern. Cell R, usually open. Face without a
sharp median ridge up its whole height (if trace of a sharp ridge ventrally then
this swelling dorsally). Antennae of varied length but not exceeding the epistomal
margin. [Not parasites of Hemiptera)

Forms with the following characters present Sirmultaneously- eyes bares anteatee
short and antennal axis at or below level of eye middle (Text-figs 29-30); gena
broad, at least as wide as length of third antennal segment and often almost as
wide as antennal length (Text-figs 29-30); no definite reclinate orbital setae;
rows of frontal setae descending to level of lunula or only just beyond; prosternum
bare; pra seta small; scutellum with three pairs of marginal setae (basals, subapicals
and strong crossed horizontal apicals, Text-fig. 70) (rarely some supernumeraries

MN

TACHINIDAE OF AUSTRALIA 29

also); abdominal T1 + 2 excavate to its hind margin; infrasquamal hairs absent;
3 with long slender L-shaped aedeagus (Text-fig. 22). [Parasites of Coleoptera]
PROSENINAE (part) (p. 41)

[Note: A few non-prosenine forms conform very closely with the characters cited.

If specimens conform but from experience do not appear to have a prosenine

facies they should be tested next in the tribal keys for Tachininae or Goniinae.

If characters appear to conform but the known host is not a beetle then pass to

couplet 8].

— Forms without such combination of characters present simultaneously, normally

failing on at least one and usually more than one of the characters cited. [Para-

sites mainly of Lepidoptera, Tapaige reed Hymenoptera, Coleoptera, Orthoptera,

Phasmatodea, Mantodea). 5 8
[Note: If the eyes are densely haired, or the prostertum is ‘ia hae; or the pra

seta is very long and strong, then pass immediately to couplet 8 without further

consideration: if the eyes are bare, the prosternum bare and the pra seta weak

then a critical consideration of the other characters cited in the first half of

couplet 7 must be made at this point. ]

8 Pre-alar seta (pra) very strong (longer and stronger than first post ia seta or the

first post dc seta) and the prosternum setulose. Propleuron bare. Second costal
sector bare ventrally. Infrasquamal hairs absent. [Parasites of Lepidoptera or
Hymenoptera Vespoidea, one genus (Pseudalsomyia) on Cerambycidae}|

GONITINAE (part) (p. 74)

— Pre-alar seta (pva) small or absent (usually not larger than first post ia seta, if larger

as in some Tachinini then prosternum bare). Prosternum bare or setulose.
Propleuron bare or haired. Second costal sector bare or haired ventrally.
Infrasquamal hairs sometimes present. [Parasites of Lepidoptera, Coleoptera,
Hymenoptera Symphyta, Orthoptera, Phasmatodea, Mantodea) z 9

9 Prosternum bare (haired in a few exceptions but then either hairing on auterior edge

of prosternum and on prosternal membrane, or arista plumose, or two strong
widely separated post ia setae and fore tarsi flattened, or epistomal margin strongly
projecting and vibrissae inserted high above level of epistomal margin, or ptero-
pleural seta absent and vein FR, setulose on most of its length). Pteropleural
seta present or absent. Epistome often strongly projecting and easily visible in
profile, the vibrissae then often high above level of epistomal margin (as in
Text-figs 41-46). [Parasites of Lepidoptera (except Apatemyia on Coleoptera) |
TACHININAE (p. 50)

— Prosternum haired or setulose but sometimes only a single hair on each side

(Text-fig. 9) (bare in a very few exceptions but then subapical scutellar setae very
strongly diverging and either apical scutellar setae absent or male with dense
hair fascicles on T5). Pteropleural seta present. Epistome not projecting and
invisible in profile or at most only rather weakly projecting, vibrissae usually
not much above level of epistomal margin (e.g. Text-fig. 49). [Parasites of
Lepidoptera, Coleoptera, Hymenoptera Symphyta, Phasmatodea, Mantodea)
GONIINAE (part) (p. 74)

[Note: It is almost impossible to separate the Tachininae from the Goniinae with
small pre-alar seta in a satisfactory way. The distinctions given in couplet 9
should help to separate specimens of the two subfamilies, but in cases of doubt
specimens should be run in the tachinine and goniine tribal keys. ]

SUBFAMILY PHASIINAE WITH KEYS TO THE TRIBES AND GENERA

This subfamily is morphologically very diversified but biologically very discrete,
for the hosts of the true phasiines are all in the Hemiptera-Heteroptera and all

30 RK. W. CROSSKEY

Tachinidae known to have hemipterous bugs as their hosts belong to the subfamily.
In the past the group has often been treated as a separate family (Phasiidae), and
certainly many of its members are very atypical looking tachinids — having widened
coloured wings and reduced or almost non-existent chaetotaxy; but these less
typical tachinids are interconnected with the more typical Tachinidae by many
forms with intermediate characters and specialists are now mainly agreed on
regarding the phasiines as a subfamily of Tachinidae. Some anomalous forms with
coleopterous hosts such as the Strongygastrini and Palpostomatini are sometimes
placed among the Phasiinae but it appears better, on the evidence so far available,
to limit the subfamily to those forms parasitic on Hemiptera.

The subfamily is moderately well represented in Australia, where all the principal
tribes, except the Gymnosomatini, occur. There is an early record of a member
of the Gymnosomatini from the Australian area, namely that of Macquart (1847 : 97
(81)) who recorded a specimen of the European species Gymnosoma rotundatum (L.)
from Tasmania, but it now seems certain that this record must be in error: Malloch
(1929a : 112) doubted the occurrence of Gymnosoma in Australia as he had never
seen the genus from this area, and no specimen has been found since to confirm
its existence there. It now appears safe to conclude positively that the
Gymnosomatini (certainly the genus Gymnosoma Meigen) are absent from Australia.

The main characteristics of the Phasiinae are as follows. Head usually without a facial
carina, sometimes with a distinct sublunular bulla between antennal bases, occasionally with
weak median ridge, only with a strong sharp keel in Eutherini; rows of frontal setae (often
weak and hair-like) descending to level of lunula or first antennal segment (rarely further);
3 without reclinate orbital setae; head sometimes holoptic in both sexes and sometimes with
greatly enlarged facets on upper parts of eyes in both sexes; eyes always bare; inner vertical
setae if present parallel or crossing; arista short pubescent; prosternum and prosternal
membrane bare; humeral callus most often with not more than two distinct setae; post ia
setae almost always o-2 (except nearly always three in Eutherini); dorsocentral setae very
varied, often much reduced; pre-alar seta present or absent, if present nearly always very
small (strong sometimes in Cylindromyiini); one or two sa setae; postalar callus with not
more than two setae; normally from one to three stf/ setae (four aberrantly) infrasquamal
hairs present or absent; usually no definite pteropleural seta; scutellum typically with two
or three pairs of marginal setae and without discal setae, sometimes only one pair of marginals
(the basals, e.g. in some Alophora), rarely four pairs (some Leucostomatini), discals usually
present in Eutherini; wing veins bare or at most with only a few minute hairs (long hairs
occasionally in Cylindromyiini) on basal node of F,,;; mid tibia usually with v submedian
seta (absent in some Phasiini); hind tibia with or without pu apical seta; suprasquamal ridge
bare; abdomen with T1 + 2 excavate only at base, sometimes virtually no excavation (except
in Eutherini where excavation reaches hind margin) ; sternites concealed or exposed; ¢ aedeagus
with non-mobile union of basiphallus and distiphallus, distiphallus without longitudinal
microstructures (‘POS’ type) (see Dugdale, 1960).

Kery To AUSTRALIAN TRIBES OF PHASIINAE

1 Wing with bend of vein M forming a gentle even curve and without trace of an M,
appendix (Text-figs 74-75). Abdomen dorsoventrally flattened (except in Savalba
with clavate abdomen) and usually devoid of strong setae. Abdominal sternites
partially or completely exposed, ventral ends of tergites not meeting in the mid
line. One supra-alar seta. One post ia seta or none. Hind tibia without pu

|

TACHINIDAE OF AUSTRALIA 31

apical seta. Scutellum with two pairs of marginal setae (sometimes only the
basal pair developed and the hind pair not differentiated). Lower calypter very
broad and rather straight or even slightly concave on its hind margin. Infra-
squamal hairs present (very minute and placed adjacent to calyptral base, care
needed to see them). Last section of vein Cu, extremely short, less than half
as long as m—cu. © without proclinate orbital setae . ; 2
— Wing with bend of vein M very abruptly angulate (usually forming a ‘sharp angle
of about 90°) and often with a short M, appendix (Text-figs 76-78). Abdomen
not dorsoventrally flattened and armed with at least a few very strong setae.
Abdominal sternites (except sometimes for St5) concealed by ventral ends of
tergites meeting in the mid line. Two supra-alar setae, a strong anterior one
and a weak posterior one (sometimes only one sa in Eutherini). Two or three
post ia setae. Hind tibia with or without pv apical seta. Scutellum with three
pairs of marginal setae in most forms (fewer sometimes in Cylindromyia and
occasional specimens of Euthera). Lower calypter slightly or strongly convex
on its posterior margin. Infrasquamal hairs absent. Last section of Cu, at
least half as long as m—cu and usually longer than this. Q with meeees orbital
setae (weak or occasionally absent in Euthera) . : 3
2 Wing with cell FR; just open or closed at the wing margin, or rich a ee short petiole
only just distinguishable (Text-fig. 74). Abdomen conspicuously elongate or
clavate, with some very weakly developed marginal setae on the posterior tergites
(these recumbent on T4 and T5, usually somewhat erect and stubby on T6).
One post ia seta present. Second costal sector haired ventrally. 92 without
externally obvious ovipositor. Eyes of both sexes widely separated, interfrontal
area very well developed and at least twice as wide as the antenna
TRICHOPODINI (p. 32)
-— Wing with cell R; closed well before the margin and with a long petiole (Text-fig. 75).
Abdomen not long and narrow, usually subovate, usually without any strong
setae (vestiture almost entirely of hairs). No post ia setae or one (if one present
this usually very weak). Second costal sector bare ventrally. 92 with obvious
projecting horny ovipositor or with large complex recurved postabdomen. Eyes
of both sexes widely separated or meeting in mid line of the frons with consequent
reduction of the interfrontal area . : PHASIINI (p. 33)
3 Palpiabsent. Posteroventral declivity of the eee feces a deep and completely
sclerotized bridge widely separating the base of the abdomen from the hind
coxae. Abdomen long and narrow (subcylindrical or slightly clavate) and with
postabdomen exposed, large and recurved. Second costal sector haired ventrally.
Hind tibia with a strong pu apical seta . : CYLINDROMYIINI (p. 36)
— Palpi present. Posteroventral declivity of the thorax membranous medially and
forming only a narrow bridge between the abdominal base and the hind coxae.
Abdomen not so shaped. Second costal sector bare ventrally. Hind tibia with
or without pv apical seta : : : 4
4 Wings colourless. Abdominal Tr ce 2 not excavate. to its hind margin. Face
without a median ridge and antennae very small (falling far short of epistomal
margin). Ocellar setae reclinate. Scutellum without discal setae. Two post ia
setae (widely spaced and anterior one subequal in size to posterior one and standing
near to transverse suture) (Text-fig. 56). Apex of 2 abdomen forcipate (Text-
fig. 93). ‘ : LEUCOSTOMATINI (p. 38)
— Wings with a bold black- brown patient (dark éolont extending along fore border
| and in two preapical cross-bands of which proximal one is wider than the distal
one, the cross-bands evanescent posteriorly; alula unusually long and narrow and
black-brown or mostly so). Face with a broad median vertical keel (sharp on
its anterior edge) and antennae exceedingly long (reaching to or beyond level of
epistomal margin). Ocellar setae proclinate. Scutellum nearly always with at

32 RK. W. CROSSKEY

least one pair of clearly differentiated preapical discal setae, these semi-recumbent.
Usually three post ia setae increasing in size from first to third, if only two present
then the anterior one remote from the transverse suture. Apex of 9 abdomen
not forcipate : : ; : : ; : ; - EUTHERINI (p. 39)

Tribe TRICHOPODINI

The Trichopodini are primarily a tropical and subtropical group and are best
represented in the Neotropics, but a few members of the tribe occur in the Ethiopian
Region, in the south-east Asian islands, in Melanesia, and in Queensland and New
South Wales. In Australia two genera occur, Savalba which is an endemic Austra-
lasian genus found from New South Wales to New Guinea, and Pentatomophaga
which is an essentially Afro-Oriental element in the Australian fauna and occurs
from Java to Queensland and New Caledonia. The African connection is shown
clearly by the fact that Pentatomophaga (although here accepted as valid, pending
revision of the Old World Trichopodini by a specialist on Phasiinae) is not really
distinct from the African trichopodine genus Bogosia Rondani (cf., for example,
Pentatomophaga bicincta de Meijere and Bogosia minor Villeneuve).

The chief characteristics of the Trichopodini are as follows. Head dichoptic, interfrontal
area wide in both sexes, eye facets not enlarged; 9 without proclinate orbital setae; face
without a median keel; ocellar setae proclinate (often very weak); antennae moderately or
very short (not nearly reaching epistome); palpi present; humeral callus with one or two
setae; ph setae nil or one; acy setae usually 0 + 1; dc setae variable, commonly I +4 2,
sometimes more; pra seta present or absent (usually absent); one sa seta; one post ia seta;
o+1or1+1 sétpl setae; infrasquamal hairs present (usually minute); scutellum with two
pairs of marginal setae, without discal setae; wings rather long and usually narrow, not
normally sexually dimorphic; second costal sector haired ventrally; basal node of R,,; bare
or with one or two very minute hairs; bend of vein M forming an open rounded obtuse curve;
cell R; open or just closed at the wing margin or with a very short petiole (less than the length
of y-m); wing membrane partially or wholly infuscate; last section of Cu, very short (less
than half as long as m-—cu); lower calypter very broad posteriorly and with outer posterior
corner rather produced, hind margin straight or slightly concave; legs weakly bristled, tibiae
not very strongly curved, hind tibia in many New World forms armed with a long fringe
of flattened black scales; hind tibia without a pd preapical seta and without pv apical seta;
hind coxae rather widely separated from abdominal base, but posteroventral declivity of
the thorax incompletely sclerotized (membranous or semi-membranous medially); abdomen
elongate and dorsoventrally flattened in most forms, sometimes slightly clavate; Ti + 2
excavate only at extreme base (virtually without any excavation); abdominal bristling very
weak, sometimes no definite setae developed (vestiture all hair like); short flattened T6 visible
from above in both sexes (usually armed with some short stubby setae on the posterior part);
© postabdomen not recurved, ovipositor very short and inconspicuous.

In both the Australian genera of Trichopodini the thoracic and leg colour is
similar: the mesonotum is velvety black or brownish black with two transverse
golden or golden-orange bands (one on the posterior half of the prescutum connecting
the notopleura and the other on the scutum immediately before the scutellum);
and the legs are yellow or orange basally (as far as the basal parts of the femora)
and black on the remainder. The head form is shown for Savalba in Text-fig. 26.

TACHINIDAE OF AUSTRALIA 33

Key To AUSTRALIAN GENERA OF TRICHOPODINI

1 Abdomen uniformly orange or yellow-orange, at most only a little darkened apically.
Sides of the abdomen subparallel or very slightly convex (Text-fig. 91). Pre-alar
seta absent. One prst dc seta. Scutum with the yellow pollinose transverse
band extending on to the postalar calli and supra-alar areas
PENTATOMOPHAGA de Meijere
— Abdomen pallid luteous yellow with a fine dark median line on the basal half and
mainly dark brown or blackish on the posterior half. Shape of abdomen distinctly
a little clavate, narrower on Tr + 2 and basal half of T3 than on succeeding
segments (Text-fig. 90). Small pra seta present. At least two prst dc setae
clearly differentiated. Scutum with the yellow pollinose transverse band more
or less confined to the prescutellar area, not extending laterally on to the postalar
calli and supra-alar areas ; ‘ : ; : . SARALBA Walker

Tribe PHASIINI

This tribe is nearly cosmopolitan, but in contrast to the Trichopodini (to which
it is undoubtedly very closely related) tends to be best represented in the temperate
and subtropical parts of the world rather than in the tropics, though many truly
tropical species are known. The tribe is represented in New Zealand, where three
species of the so-called genus Campbellia Miller occur; this genus is treated as valid
by Dugdale (1969), but appears to be indistinguishable from Mormonomyia Brauer
& Bergenstamm, a subgenus of Alophora s.l. that is widespread in Africa and
Australia.

The Australian fauna of Phasiini is comprised mainly of species (many certainly
undescribed) of the genus Alophora R.-D. This large genus occurs in both Old
and New Worlds and has been split by various authors into many so-called genera
that can at best be only very poorly defined (and are found to be largely inter-
connected by intermediate forms if the world fauna is studied). In addition to
being variously split the genus in the wide sense has often been known as Hyalomyia
(an incorrect spelling of Hyalomya), and Malloch in his various papers on the
Australian Tachinidae referred all of the described species of the complex to
Hyalomya. The names Alophora and Hyalomya were proposed by Robineau-
Desvoidy (1830) in the same work, and usage has been rather equally divided in
the taxonomic history of the group; recent works on the Eurasian and African
Phasiini have, however, mainly adopted the name Alophora for the genus, with the
name Hyalomya applied to one of the constituent subgenera where named subgenera
are recognized, and this course is here followed in accordance with the recent work
of Draber-Monko (1965). In her revision of the Palaearctic species Draber-Monko
(op. cit.) treats Alophora in a broad sense and recognizes seven segregates in the
Palaearctic fauna as named subgenera; this approach is by far the most sensible
that can be adopted in trying to deal with the difficult Alophora complex, and an
exactly comparable approach is here adopted for dealing with the Australian
fauna (which breaks into a number of moderately distinct entities taxonomically
equivalent to those treated as subgenera by Draber-Monko). At least two of the
Palaearctic subgenera, viz. Alophorella and Hyalomya, can be recognized in the

Cc

34 R. W. CROSSKEY

Australian fauna, and two other subgenera may be represented by species here
left subgenerically unplaced (these are hippobosca Paramonoy whose characters
are very close to those of the Palaearctic subgenus Phorantha Rondani, and
nigrisguama Malloch that has characters similar to those of subgenus Brumptallophora
Dupuis). Several of the Australian species, however, do not fit any of the Palaearctic
segregates, but instead are assignable to Mormonomyia Brauer & Bergenstamm,
a subgenus of Alophora s.l. until now considered to be entirely African. A careful
comparison of these Australian species with the African species (including the
type-species of Mormonomyia) has shown a complete concordance in the essential
characters, and it is here considered that the Australian species must be placed as
consubgeneric with the African ones; the subgeneric name is therefore applicable
to these Australian species. (Here it may usefully be noted that comparison of
the New Zealand genus Campbellia Miller with Mormonomyia, made whilst comparing
Australian and African species, has shown no differences that can be considered
of any nomenclatorial consequence and the name Campbellia ought almost certainly
to be placed in synonymy with Alophora (Mormonomyia).)

In addition to Alophora s.l. the Australian Phasiini includes the genera Efftayloria
and Besserioides. The former occurs also in New Guinea and is closely allied to
Alophorophasia Townsend (synonym: Kosempomyia Villeneuve), having the lower
surfaces of the femora similarly armed with rows of short stubby spinules.
Besserioides is entirely Australian on present evidence, but is evidently very closely
allied to the monotypic Ethiopian genus Bogosiella Villeneuve (from which it differs
by having the frons equally wide in both sexes and by having the posterior spiracles
of the puparium on prolonged paired bosses).

The main external features of Phasiini are as follows. Head holoptic or dichoptic, some
eye facets of both sexes enlarged in some forms; 2 without proclinate orbital setae; face
without a median keel; ocellar setae proclinate or absent; antennae usually short or very
short, if moderately long (e.g. Alophorophasia alata Townsend) nevertheless falling well short
of epistomal margin; palpi present, sometimes weak; humeral callus with o—2 setae; acy setae
usually o + 1 (hair-like); de setae variable (often hair-like), commonly 1 + 2 (sometimes 3)
post dc or only one distinct; one sa seta; one post ia seta or none; usually 1 + 1 oro + 1 sépl
setae, rarely 2 + 1; infrasquamal hairs present or absent; scutellum with one (apical) or
two (basal and apical) pairs of marginal setae, rarely weak third pair, without discal setae;
scutellum sometimes rather bullate and postscutellum often much produced posteriorly and
then visible beyond the scutellum when seen from above; wings either of normal form or
(very commonly) much produced in the anal region (especially in @) so that they are very
broad basally and short; wings often wholly or partially infuscate or yellowish or orange
basally; second costal sector bare ventrally (first sector often also bare as well); basal node
of Ry; usually with one or two very minute hairs, sometimes bare; bend of M forming an
even curve without trace of appendix; cell R; usually closed well before the wing margin
and petiolate, occasionally just open or closed at the margin (closed and petiolate in known
Australian forms); last section of Cu, very short, less than half as long as m—cu (sometimes
postero-apical corner of discal cell virtually reaching the margin of the wing); lower calypter
very broad posteriorly, hind margin nearly straight but outer posterior corner sometimes
strongly produced; legs weakly bristled in most forms or devoid of definite setae, sometimes
with a few stubby setae or combs of stubby setulae ventrally on the femora; tibiae often
strongly curved or appearing so, hind tibia without scale fringes; hind tibia without pd
preapical seta or pu apical seta; hind coxae close to or at least not very remote from the

TACHINIDAE OF AUSTRALIA 35

abdominal base, posteroventral declivity of the thorax membranous medially or more weakly
sclerotized medially than elsewhere; abdomen rounded, subovate or subconical and usually
conspicuously flattened dorsoventrally; T1 + 2 excavate only at extreme base (sometimes
virtually no excavation); abdominal bristling often weak or absent; ¢ hypopygium recurved
under abdomen, 2 ovipositor usually prominent and piercer-like; sternites fully exposed.

KEy TO AUSTRALIAN GENERA OF PHASIINI

1 Femora armed on half or more of their length with stubby black spinules on av and
pu surfaces. Pteropleuron with a moderately strong isolated pteropleural seta.
Vibrissae strong . : . EFFTAYLORIA Malloch
— Femora without such spinules, ‘yestiture of the ventral surface entirely hair-like.
Pteropleuron haired and without a pteropleural seta (at most only one of the
uppermost hairs longer and stronger than the others). Vibrissae hair-like or at
most only weakly differentiated from peristomal vestiture . ‘ 2
2 Eyes of both sexes very strongly approximated or completely meeting in the mid-
line of the frons, interfrontal area obliterated medially or if distinguishable
distinctly narrower at mid point than third antennal segment. Upper eye facets
in both sexes sometimes much enlarged compared to lowermost facets (reaching
nearly the size of an ocellus). Antennae small, third segment conspicuously less
than twice as long as second segment and falling short of epistomal margin by a
distance as great as its own length . : : . ALOPHORA Robineau-Desvoidy
— Eyes widely separated in both sexes, interfrontal area broad and at least nearly
twice the width of the third antennal segment. Eye facets of both sexes of
almost uniform small size. Antennae moderately large, third segment conspi-
cuously more than twice as long as second segment and falling short of epistomal
margin by much less than its own length. ; : : BESSERIOIDES Curran

Kry To AUSTRALIAN SUBGENERA OF ALOPHORA

1 Lower parafrontals with some fine hairing between the main frontal row of inter-
mixed fine setae and hairs and the eye. Wing with petiole of cell R; conspicuously
shorter than m—cu and less than twice as long as y-m. Mesonotum with broad
black vittae contrasting with paler greyish or sometimes partly golden pollinose
areas (either four separated vittae or the inner pair merged into a generally dark
median area). Scutellum with two pairs of marginal setae (basals and apicals) . 2

— Lower parafrontals completely bare between the main frontal row and the eye
(frontal row may consist of an admixture of fine setae and long hairs standing
very slightly out of line with each other but essentially forming one main row of
vestiture). Wing with petiole of cell R; very long, at least as long as m—cu and
more than twice as long as vm (except in A. hippobosca). Mesonotum rather
uniformly black and shining, or if partially paler pollinose then not distinctly
vittate. Scutellum with either one (basal) pair of setae or with two pairs . : 3

2 Epistome distinctly projecting, curving forwards from the facial region and the
facial profile therefore distinctly concave . . subgenus ALOPHORELLA Townsend

— Epistome not at all projecting, in the same straight plane as the face when seen in
profile (fig. 3 in Malloch, 1929a) : Alophora nigrisquama (subgenus uncertain)

3 Scutellum with two pairs of marginal setae (basals and apicals), and with uniform
fine semi-recumbent hairing on the whole upper surface. Postscutellum evenly
rounded and not, or only just, visible when fly viewed from directly above
(Text-fig. 66). @ thorax entirely without flattened lanceolate or scale-like hairs
or setae. Scutellum concolorous dark brown or black when seen from any angle,

36 R. W. CROSSKEY

slightly or strongly shining. Sublunular bulla flattened and greyish pollinose
or shining on upper part adjacent to lunula. Epistome weakly or moderately
prominent . ; 4
— Scutellum with basal setae and maelons aie genre & ifa oo weak pam bE ae
present simulating apicals then these situated in a preapical position on disc of
the scutellum) (Text-fig. 64); scutellar hairing confined to basal half or two thirds
of scutellar disc, usually the hindmost hairing rather erect and stronger than
remainder of hairing. Postscutellum very prominent, often narrowed, produced
backwards and flattened (sometimes medially slightly sulcate), always partly
visible when fly seen directly from above (Text-fig. 64). dG with some of the
hairing (sometimes also the setae) of the mesopleuron flattened and lanceolate
or broadly scale-like, similar flattened vestiture or scales often present also on
humeral callus or notopleuron [also on wing-bases and femora in some African
species]. Scutellum with posterior part ashy grey or brownish grey (conspicuously
contrasting with dark brown or black basal discal part when viewed from behind).
Sublunular bulla rounded or knob-like and brilliantly shining. Epistome enor-
mously prominent and the face seen in profile very deeply concave (Text-fig. 28)
subgenus MORMONOMYIA Brauer & Bergenstamm
4 Scutellar dorsum evenly convex. Wing with petiole of cell R, at least as long as
m—cu. Wings of both sexes clear hyaline. Claws of ¢ about equal in length to
last tarsal segment : . subgenus HYALOMYA Robineau-Desvoidy
— Scutellum distinctly flattened on pote half and with a rather sharp thin hind
margin. Wing with petiole of cell R; conspicuously shorter than m—cu. Wings
of ¢ dark brown anterobasally. iw of g enormous, obviously longer than
last tarsal segment : ‘ : . Alophora hippobosca (subgenus uncertain)

Tribe CYLINDROMYIINI

This almost world-wide group comprises some very distinctive tachinids in which,
typically, the abdomen is very elongate and subcylindrical and in which palpi
are often lacking (as in all the Australian members); in addition to these obvious
features the tribe in the strict sense is characterized by having the coxae of the
hind legs widely separated from the base of the abdomen by a completely closed
and strongly sclerotized posteroventral declivity to the thorax (normally in
Tachinidae the median part of this declivity is membranous). The limits of the
tribe are not completely clear at present as it 1s uncertain whether a number of
intermediate forms should be considered as contribal with Cylindromyia Meigen
ornot. These intermediate forms include, for example, the Oriental genus Curtocera
Macquart and Hwuttonobesseria Curran from New Zealand in which the posteroventral
thoracic declivity is closed as in Cylindromyia but in which there are strongly
developed palpi; such forms interconnect Cylindromyia and its immediate allies
with Hermya Robineau-Desvoidy, Clava Brauer & Bergenstamm, and like genera
in which there is a cylindromyiine body facies but in which the posteroventral
thoracic declivity is membranous medially (forming an ‘open’ bridge) and fully
developed palpi occur. It is not germane to consider the limits of the Cylindromyiini
on a world basis at present, but the following characteristics are noted as occurring
in all the Australian members and most of the extra-Australian members of the
Cylindromyiini sensu stricto (i.e. exclusive of forms with ‘open’ posteroventral
thoracic declivity and exclusive of forms possessing palpi).

t

TACHINIDAE OF AUSTRALIA 37

Head dichoptic, eyes widely separated in both sexes, eye facets not enlarged; 9 with two
pairs of strong proclinate orbital setae; face without a median keel or very weakly raised
medially {a definite sharp keel occurs in the Neotropical genus Polistiopsis Townsend]; ocellar
setae proclinate; antennae of varied length (much shorter than face in Australian forms, very
long and reaching epistome in some forms from elsewhere); palpi absent; humeral callus
with 2—4 setae; acy setae variable number, likewise dc setae; two sa setae; two widely separated
post ia setae (of which the anterior one is enormously strong and stands immediately behind the
presutural seta just mesad of the pre-alar seta [a very unusual chaetotactic feature in Tachinidae])
(Text-fig. 58); pra seta weak or moderately strong; one, two or three stp/ setae; infrasquamal
hairs absent; scutellum with one, two or three (most often three) pairs of strong marginal
setae; wings elongate, often partially or wholly coloured; second costal sector haired ventrally;
basal node of R,, ; with some minute hairs; bend of M forming an abrupt angle, often with
M, appendix; cell R; usually closed and petiolate (Text-fig. 76) [so in all Australian forms],
sometimes open to wing margin; last section of Cu, usually at least half as long as, sometimes
subequal to, m—cu; lower calypter evenly rounded on its hind margin; legs moderately strongly
bristled, without any scale fringes; hind tibia without fd preapical seta (only with ad and d
preapicals) and with a pu apical seta; hind coxae widely separated from abdominal base,
the posteroventral declivity of the thorax forming a deep completely sclerotized bridge;
abdomen very elongate, subcylindrical (Text-fig. 92) or subclavate (in some forms appearing
‘waisted’ in the manner of Vespoidea), with the postabdomen recurved; Tr + 2 slightly
excavate only at extreme base; abdomen with some strong setae; sternites concealed (except
for most of Sts).

All but one species of Australian Cylindromyiini belong to the genus Cylindromyna,
and the majority exactly resemble some of the common species of Cylindromyia
found in the temperate latitudes of the northern hemisphere (in fact so close is the
resemblance that if the Australian provenance were unknown it would be assumed
that specimens were either European or North American); these species have the
basal abdominal segments tawny orange or light red (usually with a dark mid line)
and the apical segments black. The single Australian cylindromyiine species that
is not a Cylindromyia was described by Bigot under the name Ocypftera tristis and
is still known only from the holotype; it is undoubtedly congeneric with Gerocyptera
marginalis (Walker) from Amboina (Ambon), type-species of Gerocyptera, and
is therefore here assigned to the genus Gerocyptera. (It is worth noting that the
type-locality of trvistis is known only as ‘Australia’, but as Bigot was sometimes at
fault in his recorded provenances there is an element of doubt whether ¢vistis is
truly Australian, which will only be resolved by future collecting: the genus
Gerocyptera occurs from the Moluccas to New Hebrides through New Guinea, and
if tvistis is indeed Australian then it probably occurs only in north Queensland.)
The head profile of G. tristis is shown in Text-fig. 25.

Some species of Gerocyptera resemble wasps of the superfamily Vespoidea and
the genus is extremely closely allied to the Neotropical genera Clinogaster Wulp
and Polistiopsis Townsend in which the resemblance to vespoid wasps is even
more perfect. (All this complex is very rare in collections and nothing is known of
the host relations.)

Kry to AUSTRALIAN GENERA OF CYLINDROMYIINI

I Lower calypter with very fine long marginal hairs in addition to the normal very
short fringe. Basal node of vein R,, ; on the ventral surface of the wing with a

38 R. W. CROSSKEY

tuft of very long strong hairs (most of these greatly exceeding the length of v—m).
Base of costa with exceedingly long hair (obviously much longer than the normal
wing fringing) extending from just distad of the basicosta for most of the way
to Sc, but especially strong immediately beyond the basicosta
GEROCYPTERA Townsend
— Lower calypter with only the usual very short marginal fringe. Basal node of
R,., ; ventrally with only a few small hairs or fine setulae, these not exceeding r—-m
in length. Base of the costa without such exceptionally produced vestiture
CYLINDROMYIA Meigen

Tribe LEUCOSTOMATINI

The small tribe Leucostomatini occurs mainly in Eurasia and North America,
but a few species of Lewcostoma are found in the Neotropical Region and in the
Ethiopian Region (principally in South Africa, but isolated specimens are known
from Sierra Leone and southern Arabia). In Australia the existence of the tribe
was known for a long time only from a single specimen collected at Sydney (in
the ANIC, Canberra collection) and identified by Malloch (1930b) as Leucostoma
simplex (Fallén) but other specimens of this species (or an extremely closely allied
one in the same genus) have recently been obtained in New South Wales, confirming
that Leucostoma occurs naturally in Australia. (The specimen named by Malloch
as L. simplex was examined during the present work and directly compared with
Fallén’s type and other European material: it appears to be correctly identified
as this Holarctic species.)

Townsend’s (1936, 1938) conception of the tribe included a strangely heterogeneous
assemblage of forms, and he even included the Australian genera Zita Curran and
Pygidia Malloch in his Leucostomatini. Whilst admitting that these genera are
hard to place reliably, it is hard to conceive of them as having any relationship with
Leucostoma. As now understood the tribe is a very homogeneous group including
forms that are all superficially very similar to Leucostoma in which the end of the
female abdomen is forcipate, and in which all the members have hemipterous
hosts. The genus Leucostoma is easily distinguished from the other genera by the
long-petiolate wing cell R; (a similarly petiolate R,; occurs in the Mexican genus
Vanderwulpella Townsend, but the female sex is unknown in this genus and the
current assignment of Vanderwulpella to the Leucostomatini is probably in error).

A noteworthy feature found in the Leucostomatini is the great development of
the lower calyptrae in the males of some forms. In the Oriental genus Calyptromyia
Villeneuve these are exceptionally large (to which feature the generic name alludes)
and dull opaque white, and in males of an undescribed leucostomatine from
Madagascar they are so enormous that they completely conceal the abdomen; in
this undescribed species the lower calyptrae are (relative to body size) without
doubt the largest known in the Diptera, and, being brilliant opaque white contrasting
with the shining black head and thorax, give the fly a most spectacular appearance.

The chief characteristics of the Leucostomatini are as follows. Head dichoptic but frons

strongly contracted dorsally in g, eye facets not enlarged; 9 with two pairs of proclinate
orbital setae (or upper pair divaricate), usually with a pair of outwardly directed prevertical

—"

TACHINIDAE OF AUSTRALIA 39

setae also; face without a median keel (at most only faintly raised on upper part); ocellar
setae reclinate or divaricate in most forms (sometimes ocellar setae almost hair-like and then
proclinate); antennae falling short of epistome; palpi present; humeral setae varied, usually
at least two (rarely one one differentiated) ; acy setae absent or o + I or 1 + I; dc setae 2 + 3
or 3 + 3; two sa setae (hind one sometimes very small); two post ia setae (very strong and
subequal in size, anterior one nearer to transverse suture than to the posterior one); pra seta
very small (rarely undifferentiated); from 1-3 stpl setae (aberrantly 4); infrasquamal hairs
absent (rarely one or two very minute hairs); scutellum with three or four pairs of marginal
setae and without discal setae (normally three marginal pairs, but additional weak fourth
pair present outside the apicals in Calyptromyia and Claivvillia Robineau-Desvoidy); wings,
normal, clear hyaline; second costal sector bare or haired below; basal node of R,,; with
one hair or strong setula (this feature very constant but a second supernumerary hair may
occur in Calyptromyia); bend of M varied, abrupt or evenly curved, sometimes with M,
appendix; cell R; open, closed at wing margin, or long-petiolate; lower calyptrae rounded,
sometimes slightly to enormously enlarged in g, often rather opaque white; last section of
Cu, at least two-thirds as long as m—cu; legs with strong setae, always without scale fringes;
hind tibia usually with three strong dorsal preapical setae (i.e. pd preapical present in addition
to ad and d), but pd preapical occasionally very weak or absent; hind tibia without pv apical
seta (except in Brullaea Robineau-Desvoidy); hind coxa and abdominal base approximated,
posteroventral declivity of the thorax membranous medially; abdomen elongate-subovate,
not flattened above, apex not recurved; Tr + 2 excavate only at base; abdomen with strong
setae; sternites concealed; end of 2 abdomen with a pair of horizontal forceps-like processes
(Text-fig. 93).

Leucostoma, the only genus known in Australia, can be recognized by the
accompanying key to phasiine tribes. The Oriental genus Calyptromyia might
just possibly occur in northern Australia though not yet found: it is distinguished
from Leucostoma by having the parafacials completely haired, four pairs of scutellar
setae, and cell Rk; open at the wing margin (instead of bare parafacials, three pairs
of scutellar setae and long-petiolate R;, as found in Leucostoma).

Tribe EUTHERINI

This tribe is currently considered to be composed of the single genus Euthera
Loew. This very distinctive genus comprises a few species from each major
zoogeographical region and an undescribed species from New Caledonia. The
tribe is very little known as most of the species are rather rare (or at least poorly
represented in museum collections). For many years the hosts remained unknown,
but it is now established that Euthera parasitizes adult Pentatomidae (Hemiptera),
a fact which confirms that the tribe is rightly placed in the Phasiinae. Before the
host relations were established the correct position of the genus was rather uncertain
and van Emden (1960 : 378, 382) put Euthera into the Minthoini because of a
resemblance to minthoines in the flattened tarsi.

Euthera was split by Townsend, who treated the Eutherini as composed of four
genera (Townsend, 1936, 1938) each confined to a different zoogeographical region
(Euthera proper in North America, Eutheropsis Townsend in the Palaearctic,
Preuthera Townsend in Africa and Macreuthera Bezzi in Australia, the last of
these originally proposed by Bezzi as a subgenus of Euthera). Separate generic

40 Re Wi. GROSSTIREY

status for these entities is no longer recognized, and certainly cannot be justified in
view of the overall homogeneity shown by eutherine species; even Townsend seems
to have thought that his penchant for splitting had been carried rather far, as he
commented (Townsend, 1938 : 212) that it ‘comes near conforming to the proverbial
genus for every species’.

In Australia the Euthera fauna contains two described species, but it is likely
that others exist, as Paramonov (1953) surmises. Paramonov (op. cit.) has noted
how newly discovered species cut across the character differences that were previously
supposed to define subgenera (or genera) within the Euthera complex, and has
emphasized the need to refrain from any further description of subdivisions within
Euthera. This is a particularly appropriate plea, as it is obvious from a study of
undescribed species in museum collections that there is a more or less complete
intergradation of characters between the species of different regions and that it is
therefore impossible to maintain any valid subgenera or genera within the complex.
Van Emden (1960) took a similar view when he discussed Euthera in the Ethiopian
Region and formally established the synonymy of Preuthera with Euthera.

The principal characteristics of Euthera and Eutherini are as follows. Head dichoptic,
eyes well separated in both sexes but frons a little contracted above in ¢; 9 usually with
proclinate orbital setae, these sometimes weakly differentiated or absent; 9 with or without
a pair of small outwardly directed prevertical setae; face with a large vertical median keel
or with a large bulbous swelling, the carination formed into a definite sharp edge medially
(Text-fig. 24); ocellar setae usually weak, proclinate or divaricate; antennae extremely long,
reaching to epistomal margin or beyond, inserted high on head; palpi present; two main
humeral setae, sometimes weak third, or third and fourth; acy setae varied, from 2 + I to
3 + 4; dc setae varied, usually at least 3 + 3, sometimes 4 prst dc or 4 or 5 post dc; pra seta
present, small; normally three post za setae, if only two then standing near to each other and
remote from transverse suture; two sa setae (hind one sometimes very weak or undifferentiated) ;
two stpl setae (I + 1); infrasquamal hairs absent; scutellum usually with three pairs of
marginal setae, but apical pair often very weak (crossed, divergent or parallel), apicals
sometimes entirely absent, discal scutellars normally present (at least one pair); wings boldly
patterned with black-brown colour (this extending along fore border and in two preapical
cross-bands which are evanescent posteriorly and of which the proximal one is much broader
than the distal one), alula unusually long and mainly black-brown; second costal sector bare
below; basal node of R,, ; bare (except in an undescribed Melanesian species in which three
or four minute hairs present); bend of M very abrupt, usually forming a right-angle or even
an acute angle, rarely with short M, appendix; cell R, long-petiolate; lower calypter rounded
posteriorly; last section of Cu, from half to two-thirds as long as m-—cu; legs with strong
setae and without scale fringes; hind tibia with two or three dorsal preapical setae, with or
without pu apical seta; hind coxa and abdominal base approximated, posteroventral declivity
of the thorax membranous medially; abdomen elongate-subovate, strongly convex dorsally,
apex not recurved; Tr + 2 with excavation reaching to its hind margin; abdomen with
strong setae; sternites concealed; no specialized ovipositor.

The most useful spot characters for the genus are provided by the combination
of variegated black-brown and hyaline wing pattern, strong median facial keel,
exceedingly long narrow antennae, and very long-petiolate wing cell R; (Text-figs
24 and 77).

It may be noted that Herting (1966 : 8) implies that the curious European
genus Redtenbacheria Schiner should be placed in the Eutherini. Such a placement

TACHINIDAE OF AUSTRALIA 41

is insufficiently substantiated at present and the diagnosis of Eutherini given
above omits Redtenbacheria from consideration.

SUBFAMILY PROSENINAE (DEXIINAE) WITH KEYS TO THE
TRIBES AND GENERA

The subfamily name Proseninae is here used in order to conform with the recently
published catalogues of the Tachinidae of North and South America, although it
is now known that it is not the oldest available name that applies to the group;
several family-group names based on included genera pre-date the use of Prosenini
(-ae) and one of these ought strictly to be used for the subfamily under the priority
requirements of the International Code of Zoological Nomenclature. It is not yet
clear which name should stand valid, and most specialists are in favour of returning
to the name Dexiinae (that used to be almost universally applied to the subfamily).
The name Dexiinae cannot, however, be validly applied to the subfamily until the
type-species of Dexia Meigen is changed by a ruling of the International Commission
on Zoological Nomenclature; such a ruling has not yet been applied for, although it
is under active consideration by interested specialists.

The subfamily is very large and cosmopolitan and its members appear exclusively
to parasitize beetles. There are very few host records for the rich Australian fauna,
but it is a reasonable guess on present evidence that all will be found to have
coleopterous hosts. Several tribes have been delimited within the subfamily by
various authors, notably Townsend, but taken as a whole the multiplicity of forms
tends to merge together with few if any breaks in the characters that permit
satisfactory tribal definitions. Even between the Rutiliini and the rest of the
Proseninae (which are conventionally treated as tribally distinct) there are few
concrete characters that absolutely serve for reliable tribal discrimination (Crosskey,
1973) of all the forms into one tribe or the other; the genus Chetogaster Macquart
could, for example, be equally well placed in either the Prosenini or the Rutiliini

(in the present work it is retained therefore in its traditional position in the latter
tribe).

As with other tachinid subfamilies it is difficult to formulate a fully satisfactory subfamiliar
definition but the main characteristics of most members of the group are as follows. Head
often with a very strong facial carina separating the antennae or with a sharp median ridge
(though carina lacking in very many forms); rows of frontal setae descending to the level of
the lunula or the first antennal segment; ¢ without reclinate orbital setae; J head often with
eyes very strongly approximated but not holoptic (except in a few Formosia species) ; uppermost
eye facets normally not enlarged (only enlarged in some Formosia); eyes bare [this is true
of almost the entire vast complex of world forms included in the Proseninae, but there are
a very few exceptions that have haired eyes: examples, Callotvoxis Aldrich and Tyreomma
Brauer and Bergenstamm from South America]; inner vertical setae when present often
convergent or crossing; prosternum bare (except in a few Rutiliuni); prosternal membrane
bare (except in some Rutiliini); humeral callus with at least two setae distinguishable (though
these may be very weak in Rutiliini); post ia setae varied, from none to four; dorsocentral
setae varied, often very reduced in Rutiliin:, pre-alar seta weak or absent; two or more
differentiated sa setae in almost all forms; postalar callus with 2-7 setae; usually two or
three sternopleural setae (one or none in some Rutiliini); infrasquamal hairs nearly always

42 R. W. CROSSKEY

absent (present at least in some species of Chetogaster); scutellum typically with three pairs
of strong marginal setae (Text-fig. 70) (including crossed or convergent apicals) but most
Rutiliini with more pairs of marginals (4-11) and apicals often reduced; mid tibia with a v
submedian seta; hind tibia with or without pv apical seta; suprasquamal ridge often haired
(Rutiliini); abdominal Tz + 2 usually excavate to its hind margin (not in Doleschallini);
abdominal sternites concealed by ventral ends of tergites meeting in mid line, at most only
slightly exposed and tergite venters slightly parted; g aedeagus long and slender and armed
with a backwardly directed spinus (epiphallus) at junctions of the basiphallus and distiphallus
(Text-fig. 22).

The curious small tribe Doleschallini, which occurs in the Oriental Region
eastwards to New Guinea and the Solomon Islands, has not been found in Australia.
It seems just possible, however, that it might be represented in northern Queensland,
and the tribe is therefore included in the following key to tribes. The Doleschallini
includes the most slender elongate forms found in the Tachinidae and the adults
are noteworthy for their habit of resting motionless (and well disguised) on the
trunks of trees (especially coconut and, in New Guinea, Casuarina) with their
exceedingly long spindly legs outstretched.

KeEy To AUSTRALIAN TRIBES OF PROSENINAE

[Note : Doleschallini is included although the tribe is not yet known from Australia. ]

1 Abdominal Tr + 2 excavate to its hind margin. Thorax membranous medially
or mainly so between bases of hind coxae and abdominal insertion, coxae and
abdomen not unusually remote from each other. Head in profile not strongly
subtriangular, as long at the epistome (or nearly as long) as at the level of the
antennal axis. Head often with a strong facial carina. Notopleuron distinctly
differentiated from the scutum by a groove or shallow depression. Body form
usually robust, but if long and slender then abdomen broadest near base or
subfusiform . : : : : : ‘ é : 2
— Abdominal Tr + 2 encavate oaly at ne bie: Thorax closed above the hind coxae
by a broad sclerotized bridge (as in Cylindromyiini), the hind coxae widely
separated from the abdominal base. Head in profile subtriangular, profrons
extraordinarily prominent and lower part of head strongly receding, head very
much longer at antennal axis than at level of the epistome. Head without facial
carina. Notopleuron not differentiated from the scutum by any depression.
Body and legs excessively long and slender, the abdomen with subparallel sides
(i.e. not distinctly fusiform or widest at base) . : : . DOLESCHALLINI
2 Postalar callus with only the normal two postalar setae (at most only a weak setula
in addition). Suprasquamal ridge and postalar wall bare. Epistome not
produced in front of vibrissal insertions (Text-figs 29-30) or only slightly so
(Text-fig. 27), usually not much visible in profile. Propleuron bare or haired.
Barette bare or sparsely haired anteriorly (if all haired then no facial carina).
Facial carina present or absent. Scutellum with three pairs of marginal setae
(some supernumerary marginals differentiated in Rutilotriva). Male hypopygium
without long strong erect setae on T7 + 8 (except in Acucera). Prosternal
membrane and prosternum totally bare . 3 : . PROSENINI (p. 43)
— Postalar callus with supernumerary strong setae, with a total of 3-6 setae (except
in Chetogaster). Either suprasquamal ridge or postalar wall usually haired.
Epistome subnasute or at least slightly and distinctly projecting in front of the
vibrissal insertions in profile (Text-fig. 31). Propleuron haired. Barette thickly

TACHINIDAE OF AUSTRALIA 43

haired along its length (except in Chetogastey in which hind part bare). Head
always with a strong facial carina. Scutellum with four or more pairs of marginal
setae (except in Chetogaster with three). Male hypopygium with long strong
erect setae on T7 + 8 (usually standing in line on each side of the tergite).
Prosternal membrane bare or haired, prosternum bare except in some forms with
haired prosternal membrane . : : c : 6 : RUTILIINI (p. 47)

Tribe PROSENINI

This tribe contains the great bulk of members of the subfamily Proseninae when
the whole world fauna is considered, but in Australia (even allowing that there
are doubtless many undescribed forms) forms only a minor proportion of the total
prosenine fauna — most of the Australian fauna being comprised by the Rutiliini.
The features that differentiate the Prosenini from the Rutiliini are shown in the
foregoing key to tribes, and the characteristics of the Prosenini need not be detailed
here (further information is, however, given in Crosskey, 1973). The Australian
Prosenini fall into two main groups, one containing forms with a heavy broad
facial carina that fully separates the antennae and with reduced palpi, and another
group (somewhat diverse in its components) containing forms without a facial
carina, or with a sharp median facial ridge, and with fully developed palpi.

The first of these groups contains the widespread Old World genus Prosena
in which the proboscis is enormously elongate and slender, and a complex of forms
closely related to Prosena in which the proboscis is much shorter and stiffer. Several
genera have been proposed among the latter forms (Senostoma, Rhynchiodexia,
Austrodexia, Macropodexia, Lasiocalypter, Lasiocalytrina) but these merge so
imperceptibly into one another — with few character breaks that are maintained
when sufficient material is examined of the complex—that it is impossible to
recognize most of them as valid genera. The oldest name applying to the complex
of forms with short proboscis, strong facial carina, and reduced palpi is Senostoma
Macquart (a name misused for many years and erroneously applied to various
rutiliines) and all the other names, except Macropodexia, are here placed as synonyms
of Senostoma; the broad genus so recognized can be identified by the accompanying
key to genera. The genus Macropodexia is superficially hardly distinguishable
from many of the species of Senostoma, but it has the propleuron thickly pale
haired and so differs from all Senostoma s.1. and from Prosena; probably Macropodexia
ought also to be placed as a synonym of Senostoma and the definition of the genus
widened, but it is here preferred to retain Macropodexia as valid until the whole
complex can be studied in more detail by an Australian worker with more material
to hand. It is of interest to note that the propleural hair is white in Macropodexia
whereas it is black in all other known Australian Prosenini with haired propleuron.

The second group contains a number of endemic Australian genera that differ
from the first group (Prosena, Senostoma complex) in having fully developed palpi
and in the conformation of the facial region of the head (either lacking a facial
carina or having a sharp ridge-like keel). The forms in this second group are rather
diversified. Most of them form an apparently natural complex in which the

44 R. W. CROSSKEY

propleuron is black haired and in which there is no facial keel, the genera Geraldia,
Acucera, Platytainia, Hobartia and Anatropomyia coming in this category; the first
three of these genera have the parafacials covered with strong black hair. This
complex seems to be uniquely Australian and has no counterpart in the fauna of
the Oriental Region, but some members closely resemble certain Neotropical forms:
Anatropomyia, for example, has the same essential characters and very closely
resembles ‘Theresia’ erecta Aldrich from southern Chile (the generic name for this
species is placed in inverted commas because the species undoubtedly is wrongly
placed in Theresia Robineau-Desvoidy but has not yet been reassigned).

Two genera here placed in the Prosenini were recently placed by Paramonov
(1968) in the Rutiliini, viz. Ola and Ruya (=Rutilotrixa). These genera (which
are so similar to each other that separate generic status is only doubtfully warranted)
have a sharp median facial keel (unlike the carina of Rutiliini or of the Prosena
group of genera) and an aggregate of characters that better places them in the
Prosenini rather than the Rutiliini, though they are to some extent intermediate.
The affinities of Ola and Rutilotrixa are uncertain, but they closely resemble some
similarly robust Prosenini from the southern Neotropical Region, especially
Psecacera Bigot.

The genus Tvichostylum is disjunct from the rest of the Australian Prosenini.
Its overall facies leaves no doubt at a glance that it belongs in the Billaea Robineau-
Desvoidy complex of genera (or so-called genera) that are a significant component
of the prosenine fauna of Africa and Eurasia. The holotype of the type-species
has the propleura totally bare (in contrast to the haired propleuron of Bullaea
s.l., Philotrichostylum Townsend, etc.) and the genus is on this account retained
as valid at present, but apart from the bare propleuron it is scarcely distinguishable
from certain species of Billaea s.l. that lack a facial carina.

The genus Heterometopia Macquart is uniquely Australian and may not truly
belong in the Prosenini (where it is here placed as an interim measure) until its
characters can be more fully studied and its hosts discovered). It is an aberrant
genus in respect of its head structure and especially difficult to place at present.
If not a prosenine then it ought probably to be placed in the Tachininae either
in the tribe Thelairini or in the tribe Leskiini. A study of the male genitalia would
probably help in determining its position.

Kry to AUSTRALIAN GENERA OF PROSENINI

t Head with a large elongate facial carina separating the antennae, the carina
convex or flattened on its anterior surface (never forming a sharp ridge) and
when seen in profile appearing abruptly marked off from the epistome by a
strong concavity (Text-fig. 27). Palpi very reduced, shorter than or at most
only as long as the third antennal segment. Two sternopleural setae (except
in Prosenina with three). Arista plumose. [Mainly slender forms in which
males have exceptionally spindly legs with attenuate, often sinuous, tibiae] : 2

— Head without a facial carina or with a rather sharp roof-like median ridge that
does not clearly separate the antennae and when seen in profile is not marked
off from the epistome by a definite concavity. Palpi well developed, obviously

TACHINIDAE OF AUSTRALIA 45

longer than the third antennal segment. Two or three sternopleural setae.
Arista micropubescent to short-plumose. [Mainly moderately or very robust
forms with legs not markedly attenuate] : : 5
Three sternopleural setae (2 + 1). Second costal = hated below: “Wing
with cell RF, closed well before the margin and distinctly petiolate (Text-fig. 79).
Proboscis very long and slender, length exceeding the height of the head (as in
Prosena). Costal spine very strong, at least twice as long as cross-vein y—m
PROSENINA Malloch
Two sternopleural setae (1 + 1). Second costal sector bare below. Wing with
cell R; open at the margin (Text-fig. 80) (except in Senostoma notatum), sometimes
narrowly. Proboscis long or short. Costal spine not very strong or undeveloped,
not longer at most than ym. . = : : 3
Propleuron haired . : : : MACROPODEXIA Pownsend
[Apart from the haired proplearos this genus does not differ from Senostoma.
The validity of the genus is therefore very doubtful]
Propleuron bare. 4
Proboscis short and straight, rather stiff, length conspicuously less hat height
of the head and usually not greater than length of the head at the epistome.
Usually two post ia setae (sometimes three, rarely only one). Abdomen with
or without discal setae. Outer part of lower calypter sometimes with long hair
on the upper surface. . ; . SENOSTOMA Macquart
Proboscis very long and slender, slightly flexible and sinuous, length conspicuously
greater than height of head. Usually one post 1a seta (occasionally two, but in
this case anterior one usually very weak). Abdomen always without discal
setae on any tergite. Lower » rage always completely bare on its upper
surface : ‘ PROSENA Le Peletier & Serville
Face with a conspicuous “root. like ridge on almost its entire height. Arista bare
or virtually so. Four or more post dc setae, usually four prst dc setae. Scutellum
with at least four distinct discal setae (in addition to the marginal setae). Inter-
mediate tergites (T3 and T4) with median discal setae, those on T3 sharply
distinguished from the very short fine recumbent hairing on either side. Abdomi-
nal T5 with strong subequal setae haphazardly arranged over most of its upper
surface. [Large and robust forms with length range 11-19 mm] : 6
Face without a conspicuous ridge on its whole height, at most with a ecitte die a
weak ridge or median swelling on the upper part immediately below the antennal
bases. Arista pubescent or plumose (pubescence sometimes very short and
imconspicuous). Three or four post dc setae, nearly always three prst dc setae.
Scutellum usually without or with two discal setae, occasional specimen with
four setae differentiated. Intermediate tergites with or without discal setae.
Abdominal T5 with strong setae usually confined to the apical row, if discal
setae present then these few and tending to form a transverse discal row. [Length
usually less than 11 mm gee of Tvichostylum rufipalpe measuring just
about 1rmm)] . 7
Propleuron haired. Scutellar margin with seven clearly ‘differentiated marginal
setae in addition to the normal three pairs. : RUTILOTRIXA Townsend
Propleuron bare. Scutellar margin on without marginal setae supernumerary
to the normal three pairs. : OLA Paramonov
[The total facies of this genus is so Sienna to ae oi Rutilotvixa that it is
questionable whether separate generic status is justified, despite the bare
propleuron. Head profile as in Text-fig. 29]
Propleuron haired. Second costal sector bare below. Parafacials bare or haired.
Two or three sternopleural setae . : : 8
Propleuron bare. Second costal sector henecel ee Dae eee ee “Thee
sternopleural setae : - : : : ; : : : : : 13

46

Io

12

13

R. W. CROSSKEY

Wing with cell R, closed well before the margin and conspicuously petiolate (the
petiole longer than v-m). Parafacials strongly haired. Three stp/ setae
GERALDIA Malloch
Wing with cell R; open to the wing margin. Parafacials bare or haired. Two or
three stpl setae . : 9
Parafacials either totally bare or italy ieaed Aneta paeseoas or (Ga Awateatanaueal
short-plumose. Two or three stp/ setae. Q palpi at most only slightly spatulate 10
Parafacials strongly haired on the upper half and with two or three minute hairs
at the ventral end adjacent to the eye, entirely bare medially. Arista long-
plumose. Three stp/ setae. 9 with extraordinarily large dorsoventrally spatu-
late palpi (coloured bright orange and conspicuously contrasting with blackish
head and body). [Moderately large form, length about Io mm, with abdomen
slightly shining blue-black or violaceous beneath the thin whitish pollinosity]

[Western Australia]. . : Undetermined gen.
[One female specimen faut eer ese seen in BMNH]

Parafacials strongly haired : : : : : - : : : : II

Parafacials bare : ‘ : é : : 4 : 3 : - - 12

Three stpl setae. 3 -+ 3 dc setae. Third antennal segment distinctly pointed on
its outer apex. Bend of vein M further from the wing margin than from m-—cu.
Humeral callus with several strong setae differentiated (usually 5-7). Palpi of 2
distinctly spatulate apically . ‘ , . ACUCERA Malloch
Two sternopleural setae. 3 + 4 dc setae. SThied teed segment evenly rounded
apically. Bend of vein M nearer to the wing margin than to m—cu. Humeral
callus with only two strong setae clearly differentiated (only at most one or two
weak hair-like setae in addition). Palpi of 2 evenly slender. Head profile as
in Text-fig.30 . : : : : : . PLATYTAINIA Macquart
Three sternopleural setae. Arista short-plumose. Epistome distinctly projecting
in front of the vibrissal insertions when seen in profile, vibrissae inserted above
the level of the epistomal margin. First antennal segment not noticeably
projecting. Antennae largely bright orange . , ANATROPOMYIA Malloch
Two sternopleural setae. Arista pubescent. Epistome not projecting when seen
in profile, vibrissae inserted about level with the epistomal margin. First
antennal segment prominently projecting. Humeral callus with three or four
setae standing more or less in line. Antennae not so coloured. HOBARTIA Malloch
Interfrontal area well developed, wider than a parafrontal in both sexes. Frons
narrower ing than 9. Arista with very long plumosity (longest hairs exceeding
the width of the third antennal segment). Hind tibia without fd preapical
seta. Wing vein R, bare. 4 without reclinate orbital setae
TRICHOSTYLUM Macquart
[This genus has very much the habitus of Billaea Robineau-Desvoidy s.l. but
has the propleuron bare. It differs from Phzlotrvichostylum Townsend by the
same character, but is otherwise exceedingly similar to this genus]
Interfrontal area very reduced or completely obliterated, in 29 obviously narrower
than a parafrontal and in ¢ eliminated by the fusion of the broad parafrontals
in the mid line of the frons. Frons as wide in gas in 9, much wider than an eye
in dorsal view. Arista with long pubescence or short plumosity (longest hairs
much shorter than width of antenna). Hind tibia with a #d preapical seta (in
addition to the normal ad and d preapicals), though this sometimes very small.
Wing with vein Ff, bare or setulose. 4 without or with one pair of reclinate
orbital setae (in latter case these standing very isolated near upper outer corners
of the wide frontal region) . : HETEROMETOPIA Macquart
[This genus, although running ont in the same couplet as Tvichostylum above,
certainly has little or no relationship with Tvichostylum and may not belong in
the Prosenini]

4

TACHINIDAE OF AUSTRALIA 47

Tribe RUTILIINI

This tribe occurs only in the Oriental and Australasian Regions, and forms a
dominant element in the tachinid fauna of Australia. A revisionary classification
of the tribe has just been published (Crosskey, 1973) and no further discussion
is needed here.

KeEy TO AUSTRALIAN GENERA OF RUTILIINI

I Suprasquamal ridge bare. Prosternal membrane bare . : : - 3 2
Suprasquamal ridge haired. Prosternal membrane bare or haired . : 5
2 Palpi minute, not longer than basal width of mentum and not longer than third
antennal segment. Buccal opening very long and narrow, at narrowest point
not wider than facial carina. Arista conspicuously short-plumose. Upper
calypter enlarged, when wings folded usually as long as lower calypter
PRODIAPHANIA Townsend
— Palpi well developed, very much longer than either the basal width of mentum or
the third antennal segment. Buccal opening wide, much broader than facial
carina. Arista pubescent (occasionally with very short plumosity). Upper
calypter normal . - a
3 Postalar callus with two strong setae (sometimes ort one — each ‘shorter and
weaker setula differentiated from hair in addition). Epistome subnasute, facial
profile deeply excavate between epistome and carina. Palpi sexually dimorphic,
distinctly clubbed or spatulate in 9 and slender in g. Infrasquamal hairs some-
times present. Scutellum normally with only three pairs of marginal setae (at
most with four, apicals included) . : - CHETOGASTER Macquart
— Postalar callus with three or more strong setae. Epistome not very strongly
prominent, facial profile usually only slightly concave between epistome and
carina. Palpi not sexually dimorphic, long and slender in both sexes (at most
only a trace of swelling at extreme tips). Infrasquamal hairs absent. Scutellum
normally with 4-11 pairs of marginal setae (including apical pair), very rare
specimens with only three pairs : 4
Apical scutellar setae inserted at a conspicuously lower level — the other sentellar
marginal setae. Postalar wall without dense hair tuft. Ventral margins of
abdominal tergites usually with rather weak vestiture, often hair-like, if moderately
strong spiniform setae present then these directed backwards as well as downwards
RUTILIA Robineau-Desvoidy (a few forms)
Apical scutellar setae inserted at the same level as the other scutellar marginal
setae. Postalar wall with dense hair tuft. Ventral margins of abdominal
tergites with very strong spiniform setae directed downwards
FORMOSIA Guéerin-Meéneville
Dorsum of thorax with four broad bold black vittae which are interrupted at the
transverse suture, appearing therefore to have eight elongate black spots (four on
prescutum and four on scutum). Palpi much shorter than mentum. Postorbits
with alternating silvery white and black spots which shift in appearance with
direction of the light. Facial carina strongly bulbous on upper part and strongly
contracted into sharp median ridge on lower part. Parafacials fully haired,
hairing reaching to a level below the lowest point of the eye and virtually continuous
with hairing on the genal dilations. [Western Australia]
CHRYSOPASTA Brauer & Bergenstamm
Dorsum of thorax without such pattern. Palpi as long as or almost as long as
mentum. Postorbits without pattern of alternating white and black areas.
Facial carina broad and flattened on anterior surface with subparallel sides, or

|

48 R. W. CROSSKEY

widened above and contracted ventrally (in latter case not normally formed
into a very narrow sharp ridge). Parafacials bare or haired, but if haired the
hairing not normally extending below the lowest point of the eye in profile and
haired area of parafacials well separated by en Ale bare area from haired
genal dilations : i : 2 : ; 6
6 Intermediate abdominal tergites achout discal setae. Facial carina usually very
broad and flattened, often slightly sulcate, with subparallel sides, at most only a
little knob-like dorsally and contracted ventrally. Abdomen without bold
black-and-white pattern (rare exceptions). [Forms often with deep median
depression in abdominal T5 or with partly metallic head]
RUTILIA Robineau-Desvoidy (most forms)
Intermediate abdominal tergites with discal setae (rarely lacking on one or both
tergites but if so then abdomen with black-and-white pattern). Facial carina
widest dorsally between junctions of second and third antennal segments and
conspicuously narrowed ventrally, not subparallel-sided and not flattened on
anterior face. Abdomen most often with bold black-and-white pattern (not
in subgenus Pavamphibolia). [Forms without wide median depression in
abdominal T5 and with the head always non-metallic] . AMPHIBOLIA Macquart

Kry To AUSTRALIAN SUBGENERA OF RUTILIA

[Note. Occasional specimens may have four postalar setae on one side and three on the
other: such specimens should be run as if four setae were present on both sides. ]

1 Parafrontals pollinose, either without metallic colour or only slightly metallic at

the vertex. Hind tibia either with well developed close-set anterodorsal fringe

or with some well developed ad or pd setae or with both fringe and sii setae.

Postalar callus with three or more setae . : : 2
— Parafrontals non-pollinose, entirely brilliant rmetalllios reo to cen violet. ” Hind!

tibia without anterodorsal fringe and without definite ad or pd setae. Postalar

callus with three setae . : ; . subgenus NEORUTILIA Malloch
2 Suprasquamal ridge bare and poetalen callus with four or five setae. Metallic

green, blue to purplish black forms with bold white pollen spots on thorax and

abdomen : ; subgenus AMENIAMIMA Crosskey
— Suprasquamal ridge ieee or = pa hed only three strong setae on postalar

callus. Colour and pattern varied, but if bold white pollinose spots present

then either only three setae on postalar callus or parafacials haired . : 3
3 Postalar callus with three strong setae. Hind tibia without a definite aritercdomsedl

fringe (occasional species with weakly developed or irregular fringe). Two or

three sternopleural setae (I + 1 or 2 + 1), anterior stp/ conspicuous amongst the

sternopleural hair. Hair of suprasquamal ridge rather short and sparse, not

noticeably crinkled and never ee on to the lower calypter (ridge bare in a

few forms) . : : : : P 3 4
— Postalar callus ae fours or more sbroue setae. Hind tibia with an anterodorsal

fringe. One sternopleural seta (o + I), at most only a very weak anterior stp/l

seta distinguishable among the hair. Hair of suprasquamal ridge long, dense

and bushy, often crinkled and often extending on to basal depression of lower

calypter (ridge always haired) : 6
4 Last abdominal tergite (T5) without a ou depression, cee Fare eaciaee ee

convex or at most with only a trace of flattening at the tip. Abdominal T3

without a transverse row of marginal setae, or if a row present then the setae not

markedly spiniform. Scutellum evenly convex on its upper surface. Supra-

squamal ridge haired. Arista long-pubescent to short-plumose. Marginal setae

TACHINIDAE OF AUSTRALIA 49

of tergite venters usually weak or hair-like and nearly completely recumbent
(not projecting noticeably downwards). ¢ sternite 5 atypical (shaped as in
text-figs 32 or 33 in Crosskey, 1973) : ‘ , : : 5
— Last abdominal tergite with a median depression, Abdominal T3 with a transverse
row of at least a few, usually many, strong erect spiniform marginal setae.
Scutellum distinctly flattened or slightly hollowed before the apex. Suprasquamal
ridge bare or haired. Arista micropubescent. Setae of inner ventral ends of
abdominal tergites rather strong and directed downwards as well as backwards.
6 sternite 5 with normal simple rounded lobes
subgenus RUTILIA Robineau-Desvoidy
3 sternite 5 very strongly acuminate on each side and provided with a pair of
submedian downwardly directed protuberances (text-fig. 33 in Crosskey, 1973).
Distal membranous part of g aedeagus exceptionally long and whip-like (about
twice as long as the sclerotized proximal part of distiphallus, text-fig. 38 in
Crosskey, 1973). Only one post ia seta (exceptionally a small second seta present
in front of main one). normally without, or with one pair of, proclinate orbital
setae. Thorax with distinct white pollinose spots or areas over mesopleuron,
sternopleuron, humeral callus and supra-alar areas of scutum
subgenus GRAPHOLOSTYLUM Macquart
— go sternite 5 without such acuminate sides and without submedian protuberances
(text-fig. 32 in Crosskey, 1973). Distal membranous part of g¢ aedeagus normal
in size, shorter than proximal sclerotized part of distiphallus (text-fig. 37 in
Crosskey, 1973). Normally two fost ia setae (one only in occasional specimens).
© normally with two pairs of proclinate orbital setae (occasionally one or none).
Thorax without distinct white pollinosity and therefore lacking bold white spots
subgenus MICRORUTILIA Townsend
6 Last abdominal tergite (T5) with a median depression and a median transverse row
of strong erect setae. Scutellum usually distinctly flattened and without distinct
preapical setae in front of the marginal row. Pteropleural hairing not developed
in front of the level of the posterior stp/ seta (text-fig. 20 in Crosskey, 1973).
Head with dark ground colour. ¢ genitalia with very large broad foliaceous
surstyli without pointed tips (text-figs 67—71 in Crosskey, 1973)
subgenus DONOVANIUS Enderlein
: — Last abdominal tergite (T5) evenly convex across its width, without median
depression or at most with only a mere trace of apicomedian hollowing: without
a transverse row of strong setae, usually only with long fine erect hairing.
: Pteropleuron haired on the anteroventral part in front of the level of the posterior
stpl seta (text-fig. 19 in Crosskey, 1973). Scutellum convex and with an irregular
row of small but definite horizontal preapical setae in front of the marginal setae.
Head usually with bright yellow ground colour. ¢ genitalia with elongate
surstyli which end in a sharp pointed tip (text-figs 72-84 in Crosskey, 1973)
subgenus CHRYSORUTILIA Townsend

KrEy To SUBGENERA OF AMPHIBOLIA

I Thorax and abdomen with bold black-and-white pattern. Apical pair of scutellar
setae approximately as strong as the other scutellar marginal setae. Lobes of ¢
sternite 5 simple and rounded on hind margin subgenus AMPHIBOLIA Macquart
— Thorax and abdomen without such pattern. Apical pair of scutellar setae much
weaker than other scutellar marginal setae (sometimes absent). Lobes of ¢
sternite 5 each with a small blunt tooth or prong on inner edge near the apex
subgenus PARAMPHIBOLIA Brauer & Bergenstamm

50 R. W. CROSSKEY

SUBFAMILY TACHININAE WITH KEYS TO THE TRIBES AND GENERA

This subfamily is to some extent an artificial assemblage of forms that cannot
readily be fitted into the other, more discretely defined, subfamilies. Some authors
split the group into two or more subfamilies, but as these have not yet been very
satisfactorily delimited on a world basis it is considered best for present purposes
to recognize the Tachininae in a broad sense. As used here the subfamily equates
almost exactly with the concept referred to as the Macquartiinae by van Emden
(1960) in his treatment of the Ethiopian Tachinidae (the nomenclature difference
arises from the fact that van Emden applied the name Yachina Meigen to the
genus that should correctly be known as Exorista Meigen and used the name
Echinomya Latreille for the genus that should correctly be called Tachina). The
usage van Emden adopted has long bedevilled the nomenclature of the Tachinidae
and is only now being gradually dropped as the correct usage under the rules of
nomenclature becomes gradually established. The subfamily here considered is
the one containing the true genus Tachina Meigen, type-genus of the Tachinidae,
and Tachininae is the correct name applying to it; the names Echinomyiinae,
Macquartiinae and Voriinae are synonyms.

The diverse nature and probable polyphyletic origin of the forms included in
the Tachininae make it difficult to define the subfamily at all precisely; to some
extent it has to be defined by the non-possession of several of the features found
in the other subfamilies. There is no single characteristic or any combination
of a few features that will absolutely serve to place any particular genus within
the Tachininae, but the following list shows the main characteristics that help
towards a definition of the subfamily on external adult characters.

Head without a facial carina separating the antennae (at most with a weakly raised median
facial ridge or with a convexity on the upper face below the antennal insertions) ; frontal setae
rows descending to the level of the first antennal segment or below; reclinate orbital setae
usually present (not in forms with strongly approximated eyes and reduced frons); inner
vertical setae often convergent or crossing; uppermost eye facets of 3 sometimes enlarged;
vibrissae nearly always well developed; arista pubescent or plumose; thoracic and abdominal
chaetotaxy often fine or reduced; prosternum usually bare; prosternal membrane bare (very
rare exceptions); humeral callus with two or more setae normally distinguishable; usually
at least three post dc setae; two or three post ia setae (less in some rare aberrant forms);
pre-alar seta almost always weaker than first post de seta, sometimes absent; normally at
least two distinguishable sa setae (one only in Palpostomatini); postalar callus with two setae
(very rarely three or only one); scutellum normally with at least two pairs of marginal setae
(one pair only in some Minthoini); sternopleural setae two or three (rarely less, very rarely
more than three); infrasquamal hairs present or absent; fore coxa sometimes haired on its
entire anterior surface; mid tibia with a ventral submedian seta (occasionally undeveloped
or minute); hind tibia with or without a pu apical seta; abdomen with T1 + 2 excavate or
not excavate to its hind margin; sternites of abdomen concealed or exposed; § hypopygium
and 9 postabdomen of varied form.

Kery To AUSTRALIAN TRIBES oF TACHININAE

1 Ocelli absent. Prosternal region greatly inflated, visible in profile. ¢ head
holoptic and with the uppermost eye facets enlarged and conspicuously bigger
than lowermost facets. Vibrissae absent : é : 2 ORMIINI (p. 57)
— Ocelli present. Prosternal region normal, no trace of inflation. g head not

TACHINIDAE OF AUSTRALIA

holoptic (although eyes sometimes very strongly approximated and frons then
very reduced), facets not obviously enlarged (except in Palpostomatini)

Epistomal region of the head prominent, visible in profile projecting in front oF
the vibrissae (sometimes head strongly subnasute) (Text-figs 41-46). Vibrissae
inserted at a level well above the epistomal margin (only slightly above in some
Leskiini). Abdominal sternites at least partially exposed between the ventral
ends of the tergites (except in Leskiini) (as in Text-fig. 20) : : :

Epistomal region of the head not prominently projecting, invisible in profile
(Text-figs 32-38). Vibrissae inserted about on a level with the epistomal margin
(undeveloped in some forms). Abdominal sternites concealed by the pte oe
ends of the tergites (except in Myiotrixini)

Abdominal sternites concealed by overlapping termite dads ree meet in aaa
venter). Scutellum with two pairs of marginal setae (basals and subapicals),
without apical setae (except in Demoticoides which has apical setae in addition).

Humeral callus usually with two or three setae : : LESKIINI (p.

Abdominal sternites at least partially exposed, ventral ends of the tergites not
meeting in the mid-venter. Scutellum with three or more pairs of marginal
setae (including a strong crossed apical pair). Humeral callus with fous or
more setae . : : : :

Hind coxa with long fine hair on the posterodorsal surface. Eyes bare. Para-

facials haired. Four post dc setae. : . TACHININI (p.

Hind coxa bare on the posterodorsal surface. Eyes haired, or if bare (Amphitropesa
and Neximyia) then parafacials also bare. Parafacials bare or haired. Three
or four post dc setae : : ; : : ; : :

Hind tibia without a pu apical seta. Two post ia setae (weak third present in
Australotachina). Postabdomen of ¢ very large, tergites 6 and 7 + 8 forming
an exposed vertical declivity at end of the abdomen. Scutellum with three
pairs of marginal setae (except Austyvalotachina with four). Bend of vein M
without an M, fold or spur-like appendix. Mid tibia without a submedian v

seta in g, with such seta in. Parafacials bare. . PARERIGONINI (p.

Hind tibia with a pv apical seta (except in Amphitropesa). Three post ia setae.
Postabdomen of ¢ not of this form, tergites 6 and 7 + 8 not externally visible
or only T7 + 8 just visible when hypopygium in situ. Scutellum with four or
more pairs of marginal setae. Bend of vein M usually with a trace of an M,
appendix or fold. Mid tibia of both sexes with a submedian v seta. Parafacials
bare or haired

Palpi very reduced, often minute or nitaally absent frecy much less than half as
long as the mentum). Parafacials haired (except in Linnaemya). Eyes densely

haired. Three post dc setae . : ; LINNAEMYINI (p.

Palpi moderately or fully developed (at least half as ; long as the mentum of the
proboscis). Parafacials bare. Eyes bare or haired. Three or four post dc setae

ERNESTIINI (p.

Sternites of the abdomen fully exposed. Vibrissae absent. Head profile as in
Text-fig. 40. Lower ends of the facial ridges expanded into a pair of broad
flattened and rather evenly haired facialia, and the epistome reduced to a long
narrow strip between the facialia connecting the oral margin directly with the
face in the same plane. Scutellum with two pairs of marginal setae (basals and

subapicals), without apical setae . : : MYIOTRIXINI (p.

Abdominal sternites concealed by overlapping ventral ends of the tergites (which
meet in the mid-line of the venter). Vibrissae present (but sometimes weakly
differentiated from the peristomal setae, especially in Palpostomatini). Head
profile not as in Text-fig. 40. Facial ridges and epistome of different form.
Scutellum usually with at least three pairs of marginal setae (two pairs only
in Palpostomatini and some Minthoini) .

67)

74)

71)

79)

55)

52

RO W. CROSSKE ¥

8 Scutellum with two pairs of marginal setae (Text-fig. 65). Prosternum with one

Io

10}

12

or two long strong downwardly directed hairs or setae on each side. Head of d

almost fully holoptic and with uppermost eye facets appreciably larger than

lowermost facets, head profile as Text-fig. 39. Supra-alar area of the scutum

with only one strong isolated seta, viz. the first sa seta, both pre-alar and second

sa setae absent (Text-fig. 54). Labellae with a pair of palpiform processes

posteriorly (usually distinct). Parafacials haired. Pallid forms with unusually

weak abdominal bristling, and very weakly developed postscutellum. [Parasites

of scarabaeoid beetles] . : : PALPOSTOMATINI (p. 53)
Scutellum with at least three pairs a eee setae (except in some Minthoini).

Prosternum bare (except in Minthoxia and Hyleorus). Head of g sometimes

with eyes strongly approximated but not virtually holoptic, uppermost and

lowermost facets not noticeably differentiated in size, head with different profile.

Supra-alar area of the scutum with at least two setae and usually three (pre-alar

seta normally differentiated but sometimes absent in Minthoini, second sa seta

present though sometimes weak). Labellae without palpiform processes.

Parafacials bare (except in Vovia with a strong downcurved seta). Not such

forms, postscutellum always strongly convex. [Parasites, where hosts known,

of Lepidoptera] . : ; : : - : : : : 9
Ocellar setae strong and as Propleuron haired. Facial ridges armed with

strong downcurved setae on their whole height. Pteropleural seta absent.

Humeral callus with four setae. Eyes haired. . CAMPYLOCHETINI (p. 60)
Ocellar setae proclinate (sometimes absent). Propleuron bare. Facial ridges

bare or with weak setulae confined to their lower halves. Pteropleural seta

present or absent. Humeral callus with fewer than four setae (except in

Nemoraeini). Eyes bare or haired 5 : 10
Wing with apical part of M (i.e. M,) and cross-vein m—cu Nemo paiee tee obkiGuer

and with the last section of vein Cu, exceptionally long (much longer than m—cu

and nearly as long as the penultimate section) (Text-figs 82-84). Head of g

with proclinate orbital and outer vertical setae (like the 2), and with frons equal

in width to that of 9. Pteropleural seta absent. Presutural intra-alar seta

present. Scutellum with long stiff erect preapical setae immediately dorsad

of the apical setae ; : VORIINI (p. 61)
Wing with M, and m—cu not ome epleee and ae ae last section of vein

Cu, short Gt most only very slightly longer than m—cu). Head of ¢ nearly

always with frons narrower than in 9 and without proclinate orbital setae (except

in Halydaia), almost always without differentiated outer vertical setae. Ptero-

pleural seta absent or present. Presutural intra-alar seta absent. Scutellum

without erect spiniform preapical setae . ‘ : ‘ 2 : : 5; II
Vein FR, setulose. Pteropleural seta absent. Scutellum with three pairs of strong

marginal setae, the apical pair crossed and horizontal and the subapical pair

very widely separated. Second costal sector haired below. Eyes bare or

virtually so. 3 + 3 dc setae ; : THELAIRINI (p. 63)
Vein R, bare. Pteropleural seta mea Searcliat not so bristled, apical setae

if present very weak. Second costal sector haired or bare below. Eyes bare

or haired. 2 + 3 or 3 + 3 dc setae : 12
Inner anterior surface of the fore coxa pose See Rance Bes dengale

long haired. Abdominal Tr + 2 excavate to its hind margin. Lower calypter

haired on the dorsal surface of its outer edge (in addition to its normal fringe-

hair). Face with a slight but distinct median ridge, visible in profile (Text-fig. 38).

Humeral callus with four or more setae. Palpi noticeably flattened dorso-

ventrally. 3-+3dcsetae . ‘ : NEMORAEINI (p. 66)
Fore coxa bare on at least most of the inner half of the anterior surface. Eyes

bare or haired. Abdominal Tr + 2 not excavate to its hind margin. Lower

TACHINIDAE OF AUSTRALIA 53

calypter bare on the entire dorsal surface, only with the usual fringe-hair. Face
without median ridge. Humeral callus with two or three setae. Palpi not so
flattened. 2+ 3dcsetae . 13
13 Abdomen with the tergites partially or ‘completely. fased on the dorsal bartace,
sutures of the mid-dorsum largely obliterated. Scutellum rotund and nearly
always with distinct preapical (dorsal) setae. Second costal sector haired
ventrally. Fore tarsus not flattened. Robust forms with luteous or orange-
red colouring and rather broad wings, and with a characteristic row of strong
ad setae along the fore tibia.. : : . GLAUROCARINI (p. 59)
— Abdomen without any fusion of the tergites and all sutures between the tergites
complete and distinct. Scutellum slightly or strongly flattened and without
preapical (discal) setae. Second costal sector bare ventrally. Fore tarsus,
especially in 9, strongly flattened. More slender or very slender forms with
mainly black coloration and long narrow wings, without a row of ad setae on
the fore tibia : : : - : - : ; . MINTHOINI (p. 64)

Tribe PALPOSTOMATINI

This curious small group, which appears exclusively to parasitize adult scarabaeoid
beetles, is very difficult to place systematically. It includes a small number of
forms with rather pallid luteous or light reddish brown colouring that are found
in Australia, Africa and the Oriental Region. It is far from clear how the group
should be defined, and it is possible that some New World forms such as Eutrixa
Coquillett and Eutrixoides Walton ought to be included within it; they appear
to be very similar to the Oriental genus Eutrixopsis Townsend, which is normally
placed in the Palpostomatini. Eutrixopsis, however, has a different conformation
to the head than typical palpostomatines, lacks definite vibrissae and has the
prosternum bare, and has a third pair of scutellar marginal setae, and it may be
a mistake to associate Eutrixopsis in the same tribe as Palpostoma; but if this
present association is correct then the palpostomatines may well have close affinity
to the ormiines and to Myiotrixa (both of which have resemblances in the head,
such as the long epistome and reduced vibrissae, to Eutrixopsis). The
Palpostomatini have not been studied in detail and in the absence of a reliable
placement they are here tentatively assigned to the Tachininae, though on certain
features they could equally well be placed in the Proseninae (with which the host
relations would certainly fit); Palpostoma itself, with its facial conformation and
bristled prosternum, even resembles some Blondeliini, and it is not entirely fanciful
to imagine some phyletic relationship to this tribe (as to some extent implied by
Mesnil’s (1966 : 882) placement (under the name Palpostomina)).

The type-genus Palpostoma is currently regarded as uniquely Australian, but
this is almost certainly only due to the fact that the group badly needs study on
a world basis. Comparison of material of the Oriental genus Hamaxia Walker
(synonym: Ochromeigenia Townsend) and the African genus Hamaxiomima Verbeke
has shown no differences of any significance from Australian material of Palpostoma,
and undoubtedly Hamaxia and Hamaxiomima ought to be treated as synonyms
of Palpostoma; it is inappropriate, however, to pursue this further in the present
work and definite synonymy is not established at this time. (The African genus

54 R. W. CROSSKEY

Hamaxioides Mesnil has not been seen but appears from description to be a true
palpostomatine distinct from Palpostoma: it differs most notably by having the
prosternum bare.)

The principal features of the Palpostomatini are as follows. Eyes bare, in g usually very
strongly approximated above and with the uppermost facets conspicuously larger than the
lowermost facets; ocelli present, often on a very strongly raised ocellar tubercle (Text-fig. 39);
inner vertical setae crossed, sometimes virtually absent in 4; proclinate orbital setae very
weak and even sometimes absent in females as well as males; face very deeply excavate and
strongly warped forwards at the epistome, but epistome vo? visible in profile; vibrissae weak
or sometimes virtually absent, level with or a little above epistomal margin, lower ends of
facial ridges often with some characteristic small stubby setulae adjacent to the vibrissae;
parafacials sparsely short haired; genal dilation weak or absent; facial ridges bare (excepting
the black setulae adjacent to vibrissae); occiput usually rather sunken; frontal setae weak
or almost hair-like, rows extending down to about the level of the first antennal segment;
arista pubescent; proboscis short, labellae usually with a posterior palpiform process (whence
the name Palpostoma); palpi well developed; prosternum usually with one or two very long
downwardly directed setae or strong hairs on each side, sometimes bare; prosternal membrane
bare; propleuron bare or haired; two strong humeral setae, sometimes minute third seta
distinguishable; o + 2 za setae in most forms, sometimes 0 + I ia; 2(3) + 3(4) dc setae; acr
setae reduced, usually 0 + 1 or 1 + 1 but variations occur; pre-alar seta absent (or at most
minute and hair-like); one sa seta only (this very strong and conspicuously isolated in the
absence of a pra or a second sa seta); 2 or 3 stpl setae; barette bare; pteropleural seta absent
or weak; scutellum usually with only two pairs of marginal setae (basals, and convergent
or crossed apicals) and without discal setae (third, lateral, pair of scutellar setae present in
Eutrixopsis); postscutellum weakly developed, sometimes almost absent; infrasquamal hairs
present or absent; fore coxa bare on inner anterior part; leg setae rather reduced, mid tibia
without or with a very small submedian v seta and with one small ad seta; hind coxa not
very remote from abdominal base, bare posterodorsally; hind tibia with two dorsal preapical
setae and without pv apical seta; tarsi not flattened; second costal sector haired ventrally;
cell R; open or short petiolate or closed at the wing margin; bend of vein M rounded or
moderately sharp and appendiculate; veins bare above except for fine hairs on R,, ; that
reach part way to, or fully as far as, yv-m; last section of Cu, as long as m—cu; distance on
vein M from bend to m-—cu usually as long as or longer than distance from v—m to m-—cu;
lower calypter very strongly diverging from scutellum, evenly rounded on its hind margin;
abdomen with weak bristling and Ti + 2 not excavate to its hind margin; T1 + 2 without
median marginal setae; sternites concealed.

The Australian fauna includes two genera in addition to Palpostoma, viz.
Apalpostoma and Eustacomyia; these are very little known genera, not recorded
since the original descriptions of their included species, but are here accepted as
valid genera distinct from Palpostoma. The Oriental genus Eutrixopsis is included
in the following key, although as already pointed out there is some doubt as to
whether it truly belongs in the Palpostomatini.

KeEy To AUSTRALIAN GENERA OF PALPOSTOMATINI

I Prosternum bare. No definite vibrissae or vibrissal angle. One post ta seta.
[Oriental Region including Japan] . - c . EUTRIXOPSIS Townsend
— Prosternum with one or more long seta or Safe on each side directed downwards.
Vibrissae distinctly developed and head with definite vibrissal angle. Two
post ia setae. [Australia] é 5 ; : : ¢ : . 6 3 2
2 Propleuron bare

TACHINIDAE OF AUSTRALIA 55

— Propleuron haired . : . EUSTACOMYIA Malloch
3 Wing with cell R; open to te wing eet oil bend of vein M sharp and provided
with a definite short M, appendix . : : . APALPOSTOMA Malloch
— Wing with cell R,; closed at or before the wing margin (usually short petiolate) and
bend of vein M without trace of appendix . PALPOSTOMA Robineau-Desvoidy
{The extralimital genera Hamaxia and Hamaxiomima differ from Palpostoma
only in having cell R,; narrowly open at the wing margin, like Apalpostoma. The
closure of cell R, probably should not be considered sufficient for generic separa-
tion and Hamaxia and Hamaxiomima would be best placed as synonyms of
Palpostoma on a redefined basis. Similarly Apalpostoma should probably be
merged with Palpostoma if the presence of a short M, appendix at the bend of
M is the only remaining distinction]

Tribe MYIOTRIXINI

Townsend (1936) erected this tribe for the single genus Myiotrvixa Brauer &
Bergenstamm. This-is a very aberrant tachinid that is still known only from the
male holotype of WM. prosopina in the Vienna Museum. The affinities of this species
are very obscure and it is therefore appropriate to retain it in its own tribe until
its relationships can be determined. The general appearance of the head with
its very reduced antennae, lack of vibrissae and broadly dilated flattened setulose
facialia is reminiscent of certain ormiines or ormiine-like forms and there may be
some relationship between Myzotvixa and these forms, or perhaps with the
Palpostomatini; any suggestions on the affinities are mere guesswork, however,
in the absence of any knowledge of the female, the early stages or the host relations
(the overall appearance conveys the impression that Myiotrixa might parasitize
beetles but there is no evidence for this at present); possibly the Myiotrixini should
be placed in Proseninae.

As Myvotrixa has been rather poorly described up to now, and not previously
figured, the opportunity is here taken to provide a figure (Text-fig. 40) of the curious
head and a description of the principal characters shown by the type-specimen
(the thorax of the holotype is badly damaged where it is pierced by the large
mounting pin but the specimen is otherwise in fair condition and most of the
characters can be made out without difficulty: it appears that the specimen was
still rather teneral when collected as the lunula and antennal bases are still partially
retracted).

Head profile as in Text-fig. 40; eyes rather strongly oblique, with very short sparse hairing;
vertex about twice as wide as the antennae, slightly sunken around the ocellar triangle; ocelli
present, ocellar setae absent; head drawn down anteroventrally so that in profile it forms
a rather sharp angle at the epistomal margin; epistome completely invisible in profile,
completely flat and equibroad with the face, very sharply differentiated from facialia; facial
ridges evanescent above but expanded and flattened below and reaching to anteroventral
corners of the head; genal dilation well developed, obliquely reaching to anteroventral corner
of the head; gena about one-fifth of eye height; parafacials very narrow, at mid point much
narrower than the antennae; interfrontal area well developed, about twice as wide as a
parafrontal. Inner vertical setae convergent, outer vertical setae present and strong; frontal
setae in about eight pairs, inclinate and crossed, rows not reaching much below level of the
lunula; two pairs of long fine erect orbital setae that are scarcely differentiated from the

56 R. W. CROSSKEY

frontal rows; no proclinate orbital setae; interfrontal area sparsely haired; parafrontals
haired; parafacials haired on most of their height (only bare at lower ends) with similar dark
hairing to that on the parafrontals; vibrissae absent; facialia with a row of long rather weak
setae along the front inner edge (when head seen in profile the row extending from the level
of the apex of the antenna to the anteroventral corner of the head) and elsewhere covered
with long strong hairing; setulae of postocular row very weak; occiput flat above; postbuccal
regions with fine dark hairing, postgenal and mid occipital regions with yellowish hair.
Antennae inserted about level with eye middle or slightly above, head about as long at
antennal axis as at the epistome when seen in profile; antennae exceptionally small, third
segment almost orbicular and only a little longer than second segment; antennae falling
short of epistome by a distance very much greater than their own length; arista pubescent,
thickened only at its extreme base; proboscis very short, palpi large and flattened and
unusually long haired. Thorax. Notopleuron rather strongly triangular, with usual two
notopleural setae. Prosternum, prosternal membrane and propleuron bare. Propleural
seta minute, one strong prostigmatic seta. Anterior spiracle without hair fringe around the
opening, sclerotized area of the spiracle large, slightly convex and sparsely long haired.
Humeral callus with two strong setae. Pteropleural seta absent, sternopleural setae I + I.
Three strong hypopleural setae; pleurotergite bare. [Mediotergite dirty or damaged,
condition of infrasquamal hairs unknown.] SBarette sparsely haired anteriorly. [Chaetotaxy
of mesonotum uncertain owing to extensive damage but apparently as follows: 1 + 3 za setae
(of which prst¢ and first post ia hair-like); probably 2 + 3 dc; at least 1 + 2 acr setae but
probably more; small pre-alar seta.] Scutellum with only two pairs of marginal setae (basals
and subapicals), no lateral or apical setae or definite discal setae; subapical setae slightly
divergent and rather weak, distance between their bases subequal to that between a subapical
seta and its corresponding basal seta; hairing of scutellar disc rather weak but long and
semi-erect. Postscutellum very strongly developed. Legs. Rather short and with very
reduced chaetotaxy. Fore coxa bare on inner anterior surface; fore tibia with two minute
pu setae. Mid tibia without a submedian v seta, with one very small ad seta on apical half
and with two or three minute p setae. Hind coxae rather close to abdominal base, bare
on posterodorsal edge; hind tibia with several very small ad and pd setae or setulae, with
one small av seta, without definite pu apical seta, and without the normal d preapical seta
(the dorsal preapical setae represented only by the ad preapical). Wings. Narrow, very
faintly smoky brownish, especially anteriorly. Vein R,,; with minute setulae extending
from node about half-way to v-m, wing veins otherwise bare. Second costal sector bare
below. Cell R,; well open at wing margin. Bend of vein M evenly and gently rounded, no
trace of appendix; distance on M from bend to m-—cu subequal to that between 7m and m-—cu;
m-—cu straight, apical section of Cu, very short and only about half as long as m—cu. Second
costal sector short (length not exceeding that of m-—cu). Sixth vein stopping well short of
wing margin. Calyptrae normal, lower one moderately broad and bare on dorsal surface.
Abdomen. Short subovate with Tr + 2 not excavate to its hind margin; each visible tergite
subequal in length. Sternites exposed, isolated in wide membranous area of abdominal mid
venter. Abdomen completely devoid of bristles, vestiture composed solely of short fine
recumbent hairing (the hairing only slightly longer at tergite margins than elsewhere and
nowhere erect). All sutures between tergites normal, no trace of fusion. ¢ hypopygium
small, no noteworthy external characters [not examined at this time as holotype unique:
later examination of male genitalia may well shed light on the affinities]. Colovation. Almost
entirely blackish brown; antennae, lower end of interfrontal area and area of head between
genal dilation and facialia paler reddish brown; abdomen covered with pale yellowish pollinosity
that is thickest on basal parts of the tergites, the tergites therefore appearing darkest towards
hind margins and also medially but appearance slightly shifting with light direction. Body
hair black or brownish black except for some yellowish hair posteriorly on the head. Measure-
ments. Body length about 6-0 mm, wing length 5-3 mm.

The holotype bears three handwritten ink labels that read ‘56’, ‘Thorey 1864

TACHINIDAE OF AUSTRALIA 57

Austra. sept’ and ‘N.G. [=new genus] ad Oestrophasia N. Holl.’, and Brauer &
Bergenstamm’s determination label that reads ‘prosopina det. B. B.’ (specific name
handwritten, remainder printed).

Tribe ORMIINI

This small tribe occurs mainly in the tropical and subtropical parts of the world,
and contains some of the most aberrant forms to be found in the Tachinidae.
Several genera have the chaetotaxy very reduced and have a robust form and curious
head facies reminiscent of the Oestroidea. Biologically the group is particularly
interesting because the first-stage larva is the most perfectly formed planidium
occurring in the Diptera (though Glawrocara has a similar planidium: see under
Glaurocarini) and the hosts appear always to be nocturnally active Orthoptera
(crickets and Tettigoniidae s.l.). Many forms have the pallid coloration often
associated with nocturnally active flies.

At present the Ormiini are uniquely characterized by having the prosternal
region of the thorax greatly inflated (usually more so in females than males) and
visible when the fly is seen in profile. The function of this oddly modified prosternum
is unknown; such a feature occurs in no other Tachinidae. Another very unusual
character among Tachinidae that occurs in many Ormiini is the reduction of the
ocelli, some genera having the ocelli vestigial or totally wanting; all the ormiines
so far known in the Australasian fauna and in the African fauna are without ocelli.
In the Oriental fauna the genera Homotrixa Villeneuve and Xanthooestrus Villeneuve
possess ocelli, but in these genera the prosternal membrane is only moderately
inflated (Homotrixa) or not noticeably inflated at all (Xanthooestrus) and it is not
certain that the genera truly belong in the Ormiini.

The existence of forms such as Xanthooestrus, which has an entirely ormiine
facies but has a normal prosternum, and of forms such as Glaurocara, which has
a different morphological appearance yet has a perfect planidium larva and host
relations like the Ormiini, poses the question of whether the Ormiini ought to be
redefined on a much wider basis than at present. It appears now that a definition
of the Ormiini to include only those forms with inflated prosternal membrane is
too restrictive, and that several forms placed outside the group at present should
be brought into it. At present, however, insufficient is known of these generally
rather rare forms to permit any worthwhile re-evaluation of the Ormiini on a world
basis, and the Ormiini is here accepted in its usual confined sense with the
Glaurocarini recognized as a separate (rather intangibly defined) tribe.

The main characteristics of the Ormiini are as follows. Eyes very large, bare, facets
conspicuously enlarged in most males; head of ¢ usually holoptic or almost so, ¢ frons
therefore virtually absent; ocelli present or absent, when present often strongly raised in g;
inner vertical setae absent in male, usually present and converging or crossed in 9; proclinate
orbital setae absent in g, weak in 9; parafacials bare; epistome not projecting, invisible in
profile, often formed into a receding strip which curves for a long distance between a reduced
face and the oral cavity; oral cavity, proboscis and palpi often exceptionally reduced; vibrissae
present or absent, when absent lower ends of facialia usually formed into broad haired areas
on each side of the epistomal strip; genal dilation weak or virtually absent; occipital region

58 Re. Wi. CROSSKEY

flat; frontal setae reduced or hair-like, rows reaching about to level of first antennal segment;
antennae small or minute, third segment rounded apically and usually not longer than second
segment, ends of antennae falling short of margin of oral cavity by a distance at least equal
to length of third antennal segment and usually by a distance much greater than their own
length; arista bare to short plumose; prosternal region inflated, prosternum and membrane
bare; propleuron bare; propleural and prostigmatic setae often hair-like; humeral callus with
two strong setae in most forms; mesonotal chaetotaxy varied, often much reduced (Text-fig. 55),
normally 2 + 3 de setae and o + I or o + 2 ia setae (rarely a minute third post ia or no ia
setae at all); pva seta small or very small, either one or two sa setae; usually two stp/ setae,
sometimes only one; pteropleural seta absent or present (if present usually weak) ; infrasquamal
hairs absent; scutellum usually with three pairs of marginal setae, the apicals usually crossed;
fore coxa usually bare on much of inner anterior surface; mid tibia without submedian v seta
or with a small v seta; hind coxae bare posterodorsally, not remote from abdomen; hind
tibia with small pu apical seta and two dorsal preapicals; wing with second costal sector
usually rather long and haired ventrally; costal margin sometimes (in New World forms only)
with callus-like dilation between apices of R, and R,,, in g; cell R; open or closed; bend of
vein M very sharp, usually about a right-angle, usually with short M, stump-like appendix;
distance from bend of M to m-—cu shorter than or subequal to m—cu/y-m; upper surface of
wing veins totally bare or with a few minute hairs only on basal node of R,,;; lower calypter
broad, inner posterior angle close to scutellum; abdomen rotund, usually with much reduced
chaetotaxy, often without any definite bristles; Tr + 2 excavate to hind margin; sutures
of abdomen distinct; intermediate abdominal tergites always without discal setae; sternites
widely exposed; 9 terminalia modified for deposition of planidia.

In Australia the Ormiini were unknown until Paramonov (1955) described two
species from Queensland and assigned them to the supposedly new genus Ovmiominda
Paramonoy. In an earlier paper (Crosskey, 19660) it was shown that Ormiominda
cannot be distinguished from the widespread Old World ormiine genus Therobia,
and the opinion given earlier that Ovmiominda is a synonym of Therobia is here
upheld as undoubtedly correct. Therobia is the only genus of the tribe yet known
in Australia, but it is possible that Phasioormia Townsend occurs in north Queensland;
the latter genus occurs from Burma to Papua and may ultimately be found in
Australia (hence its inclusion in the key that follows). Finally it may be noted
here that Paramonov’s species of Thervobia from Queensland seem rather doubtfully
distinct, and it is also uncertain whether either or both are really separate species
from T. braueri (Kertesz) from New Guinea, T. vesiculifera Bezzi from Fiji, or
T. insularis Séguy from New Caledonia (some or all of which may well be synonymous
with each other).

Kery To AUSTRALIAN GENERA OF ORMIINI

1 Epistome forming a long narrow strip running from the oral cavity to the antennal
apices and flanked on each side by broad flattened and rather evenly haired
facialia. No vibrissae. Antennae minute, length much less than a quarter of
eye-height, apices separated from the oral cavity by a distance very much greater
than their own length. Oral cavity extremely reduced and subcircular. Frons
of 9 usually noticeably tapering towards the ventralend . . THEROBIA Brauer

— Epistome normal, without such form. Vibrissae distinct. Antennae not excep-
tionally small, length greater than a quarter of eye-height, apices separated from
oral cavity by a distance less than their own length. Oral cavity not exceptionally
reduced, normally obviously longer than its width. Frons of 2 with parallel sides

PHASIOORMIA Townsend

TACHINIDAE OF AUSTRALIA 59

Tribe GLAUROCARINI

This is a very small tribe that is confined to the Old World tropics and subtropics
and contains only the two genera Glaurocara Thomson and Doddiana Curran;
other supposed Old World genera have been described (Oestrocharis Villeneuve,
Semisuturia Malloch and Oestrocara Townsend) but these are now treated as
synonyms (Crosskey, 1962). Townsend (1936) placed several New World genera
in the tribe but none of these have the fused abdominal tergites of true Glaurocarini
and evidently do not belong. Doddiana occurs from Java to Queensland and
New South Wales, and Glaurocara occurs in Malaya, the south-east Asian archipelago,
the Ethiopian Region and Mauritius; Mauritius is the type-locality of G. flava
(type-species of Glaurocara) and is also a locality into which Doddiana mellea
(Wiedemann) has been unsuccessfully introduced from Java for control of sugar-cane
moth borer.

The only host recorded for the genus Doddiana is the sugar-cane moth borer
Chilo sacchariphagus (Bojer) (Lepidoptera: Pyralidae), which is apparently attacked
by Doddiana mellea in Java. This is a curious record, as Doddiana is without
doubt very closely allied to Glaurocara and the type-species of this genus, viz.
G. flava Thomson from Africa, is a parasite of nocturnally active tettigoniid
Orthoptera (Crosskey, 19655). It is odd that species of two such similar genera
as Doddiana and Glaurocara should attack hosts as different as a pyralid and a
bush-cricket. It seems probable, however, that Orthoptera are the normal hosts
of the Glaurocarini, as Glaurocara first-stage larva is a perfect planidium closely
similar to that of the Ormiini and the ormiines (so far as is known) are exclusively
parasites of Grylloidea and Tettigonioidea. Glawrocara also resembles the Ormiini
in being nocturnally active and in having the pallid luteous yellow coloration
characteristic of night-flying Diptera, and its female reproductive system is of
Townsend’s type XXXII which is elsewhere found only in the Ormiini. On these
grounds it appears that the Glaurocarini are closely allied to the Ormiini, although
they differ obviously from the latter tribe by having the sutures between the
abdominal segments largely fused above and in having a normal prosternum (without
trace of the inflation of the prosternal region as occurs in Ormiini).

The chief characteristics of the tribe are as follows. Head unusually small for body size;
eyes usually with some very minute sparse hairing (appearing bare at first glance); ¢ head
with eyes strongly approximated and frons reduced but not holoptic; ocelli present, usually
rather raised; inner vertical setae present in both sexes, usually convergent or crossed; ¢
without proclinate orbital setae, 9 with two pairs (very strong); 9 with one pair of reclinate
orbital setae; parafacials bare; frontal setae moderately strong, rows reaching to a level with
first antennal segment; epistome flat, invisible in profile, face rather flat; oral cavity, proboscis
and palpi very small; vibrissae present, inserted about level with or just above epistomal
margin; facial ridges with a few setulae above vibrissae, otherwise bare; genal dilation weak,
only reaching at most half way forwards on the genal region, usually bearing some rather
strong setae or setulae (Text-fig. 34); occiput moderately flat above; antennae small (length
about half the eye height), inserted about on a level with eye middle; arista with short or
long pubescence; prosternum not inflated, bare, prosternal membrane bare; propleuron bare;
prostigmatic and propleural setae very strong; humeral callus with two strong setae (sometimes

a weak third seta present mesad of main two); mesonotal chaetotaxy usually as in Text-fig. 57;
o + 2 or o + 3 ia setae; 2 + 3 dc setae; acy setae variable; pra seta weak or absent, second

60 Re Wi (GROSSE MS

sa seta present but very weak; pteropleural seta present; 2 stp/ setae; infrasquamal hairs
absent; scutellar setae very variable, but always at least three pairs of marginals, subapical
pair subparallel or weakly divergent and inserted close together, discal scutellar setae almost
always present, scutellum unusually rotund; fore coxa bare on inner anterior half, fore tibia
with a row of strong ad setae or setulae along its length (usually also with some small p setae
in addition to the normal pv setae), mid tibia with a submedian v seta; hind coxae not remote
from abdominal base, bare posterodorsally; hind tibia without pv apical seta, with two dorsal
preapical setae; wings relatively large compared to body size; second costal sector haired
below; cell R; narrowly open at wing margin; bend of vein M abrupt, with or without M,
appendix; distance from bend of M to m—cu less than that between v—-m and m-—cu; last section
of Cu, shorter than m—cu (Text-fig. 81); wing veins bare above except for some minute setulae
on basal node of It4,;; lower calypter moderately broad, inner posterior angle near to
scutellum; abdomen rotund, dorsally with the sutures between the tergites mainly obliterated
by tergite fusion; Tr + 2 excavate only basally; sternites concealed.

KrEy TO GENERA OF GLAUROCARINI

I Bend of vein M very abrupt and with a distinct M, appendix. Tip of the scutellum
with a pair of small fine straight apical setae between the strong close-set subapical
setae (if rarely there are no distinct apicals then tip of scutellum with at least
some hairs between the subapical setae) . : . GLAUROCARA Thomson
— Bend of vein M moderately abrupt but “ayegt trace of M, appendix. Tip of
scutellum without apical setae and usually totally bare Poe the bases of the
strong close-set subapical setae : : : : : . DODDIANA Curran

Tribe CAMPYLOCHETINI

Townsend (1936) erected this tribe for a small number of genera found in each
of the major zoogeographical regions and apparently closely allied to the Voriini.
The group is easily recognized by having the strong ocellar setae directed backwards
(a very unusual character found elsewhere in Australian Tachinidae only in the
Goniini and in Leucostoma) and simultaneously having the propleuron haired and the
facial ridges strongly bristled. Until now the tribe has not been reported from
Australia, but recent collecting has shown the presence of some undetermined (probably
new) species in Australia and Tasmania that clearly belong in the genus Elpe
Robineau-Desvoidy; this is the only genus so far discovered in Australasia. The
genus Elpe is widespread in the Palaearctic, Ethiopian and Oriental Regions but
apparently does not occur in the New World. Mesnil in various publications has
referred several species of the genus to Frivaldskia Schiner (=Fallemia Meigen,
preoccupied), but the true Frivaldskia (type-species F’. longicornis (Fallén)) has the
parafacials partially haired and cell R; closed and appears to be generically distinct
from Elpe (van Emden, 1960 : 351). (Here it may be noted in passing that
Frivaldskia is the correct original spelling and that the various alternatives such as
Frivaldszkia and Frivaldzkia that have currency in the literature are incorrect
subsequent spellings.)

Van Emden (1960) placed Elpe as a subgenus of Campylocheta Rondani, but
the latter has a different facies from Elpe and has three (instead of two) stpl setae
and extensively hairy parafacials (instead of bare parafacials as in Elpe). It

TACHINIDAE OF AUSTRALIA 61

appears best to treat Elpe and Campylocheta as separate genera, but Frivaldskia
ought probably to be treated as a synonym of Campylocheta.

The principal characters of the tribe are as follows. Eyes haired; facial ridges strongly
setose (Text-fig. 33); face deeply sunken but epistome not projecting; vibrissae level with
epistomal margin; antennal axis well above level of eye middle; head receding below, much
shorter at vibrissal axis than at antennal axis; ocellar setae very strong and directed outwards
and backwards; inner vertical setae parallel; parafacials bare, or haired on upper half only;
proboscis very short, palpi well developed; prosternum and prosternal membrane bare;
propleuron pale haired; humeral setae four (basal row of three and one set forwards) ; 2-3 + 3
dc setae; pre-alar seta very small; 1 + 3 ia setae; two or three stp/ setae; scutellum with three
pairs of very strong marginal setae (basals, subapicals and horizontal crossed apicals), subapicals
extremely wide apart, occasionally a weak fourth pair of setae present between the subapicals
and apicals; scutellum without definite discal setae; pteropleural seta absent; infrasquamal
hairs present or absent; fore coxa bare on inner anterior surface; mid tibia with a v seta;
hind tibia without pv apical seta, usually with three strong d preapicals; hind coxa bare
posterodorsally; wing with several strong setulae dorsally on node, veins usually otherwise
bare above (some American forms with /, setulose); A; open or closed in wing margin
(occasionally very short-petiolate); bend of vein M forming very obtuse angle, without
appendix, remote from wing margin; distance on vein M from bend to m-—cu at least equal
to and usually greater than that from vm to m-—cu; apical section of Cu, longer than m-—cu;
second costal sector very short and haired ventrally; calyptrae normal; abdomen with Tr + 2
excavate to, or almost to, its hind margin; Tr + 2 without median marginal setae; intermediate
abdominal tergites with discal setae; T5 with a complete transverse row of very strong erect
discal setae around the upper surface; sternites concealed; male hypopygium very conspicuous,
T7 + 8 at least partially exposed, cerci enormous and curved forwards (slightly hook-like)
in profile.

Finally here it should be noted that the spellings Campylochaeta and
Campylochaetini in the literature are erroneous, as Campylocheta Rondani is the
correct original spelling in nomenclature.

Tribe VORIINI

Most of the members of this tribe have a very characteristic head facies and
wing venation and it is usually possible to recognize the voriines easily once one
of the genera has been seen in a named collection; often one glance at the wing
is sufficient for reliable tribal placement. The notable features of the head and
wing alluded to are listed in detail later. The tribe is nearly cosmopolitan, and
includes four genera in New Zealand; in this tribe there is therefore, perhaps, some
faunal connection between Australia and New Zealand. Three genera are known
from Australia, all of which occur also outside the area: Voria occurs widely in the
New World and throughout most of the Old World, A/frovoria occurs in Africa,
India and Western Australia, and Hyleorus occurs in New Guinea as well as Australia.
When the Voriini are studied in more detail it will almost certainly be concluded
that Hyleorus is widespread also in the Ethiopian Region and Eurasia (including
Japan) as there appears to be little doubt that the genus Stezmiomyia Townsend
(synonyms Neuroplagia Townsend and A froplagia Curran) is identical with Hyleorus;
comparison of material of several species of Stewmiomyia (including elata Meigen,
the type-species) with Hyleorus from New Guinea and Australia has shown no

62 kK. OW. CROSSIKE Y

differences that could fairly be interpreted as of generic significance, but it is outside
the scope of this work, to consider this further at present (as Hyleorus has priority
and will stand as the valid name in any case) and definite synonymy is not established
at present.

The main characteristics of the Voriini are as follows. Head not noticeably sexually
dimorphic, both sexes with outer vertical setae and proclinate orbital setae; eyes bare or haired;
ocellar setae divaricate-proclinate or absent; inner vertical setae subparallel; facial ridges
bare, or weakly setose for half their height; parafacials usually with hairs or strong down-
curved setae on part of their height, sometimes bare; face rather flat, epistome not projecting
(invisible in profile); vibrissae level with epistomal margin; upper occiput without black
setulae; antennal axis about level with or slightly above eye middle; proboscis short, palpi
well developed; antennae often with second segment unusually long in relation to third, arista
thickened on at least half its length, often with second basal segment very elongate; prosternum
bare or haired, prosternal membrane bare, propleuron bare (haired in a few rare exceptions) ;
humeral callus with two strong setae and usually a third smaller seta mesad of main two and
one small seta set forwards; mesonotal chaetotaxy typically as Text-fig. 59; I + 3 7a setae;
2 or 3 + 3 dc setae (very rarely four post dc); pre-alar seta very small; three sternopleural
setae (very rarely only two); pteropleural seta either absent or enormously strong; infrasquamal
hairs present or absent; scutellum with three pairs of strong marginal setae (basals, widely
spaced subapicals and crossed horizontal apicals), and nearly always with a pair of long strong
stiff erect preapical setae immediately above the apicals (sometimes some additional long
stiff erect setae further forwards on the disc of the scutellum in addition); fore coxae usually
haired on most of its inner anterior surface; fore tibia typically with a series of definite ad
setae or setulae along its length; mid tibia with a v seta; hind coxa bare posterodorsally; hind
tibia with or without small pu apical seta and with two d preapicals; wing extensively setulose
dorsally on the veins (R,,; setulose on much of its length, R, and Cu, also often setulose) ;
bend of vein M strongly angulate and remote from wing margin, M continued towards the
wing margin by a long or often very long /, spur vein or by a darkened fold (Text-figs 82-84) ;
last section of vein Cu, much longer than m—cu and nearly as long as or even longer than
penultimate section; M, and m-—cu usually exceptionally oblique (some New Zealand forms
with m-—cu non-oblique); cell R; usually open at wing margin, sometimes closed in’ margin
or long-petiolate; second costal sector short or very short, bare or haired ventrally; abdominal
Tr + 2 excavate to its hind margin or almost so; sternites concealed; ¢ hypopygium with
exceedingly long slender curled aedeagus.

KeEy To AUSTRALIAN GENERA OF VORIINI

I Parafacial bare, without either hairs or strong downcurved setae. Arista thickened

on more than half its length, second aristal segment at least twice and usually
more than twice as long as its greatest breadth - ; 2

— Parafacial with a very strong downcurved seta inserted at the ape end immediageie

below the lowermost foaral seta (Text-fig. 37). Arista thickened only on its

basal half, second aristal segment not longer than wide or only inconspicuously so
VORIA Robineau-Desvoidy

2 Prosternum haired (sometimes only a single hair on each side). Eyes obviously

hairy. MM, spur vein exceedingly long, usually much longer than the section of

vein M between m-—cu and the bend (Text-fig. 84) [slightly shorter in an unde-

scribed species from Philippines]. Both intermediate abdominal tergites with

erect discal setae. Facial ridges setulose on half their height (in profile no gap

between the uppermost setula on the facial ridge and the lowermost frontal seta)
HYLEORUS Aldrich

nl

TACHINIDAE OF AUSTRALIA 63

[The Eurasian and African genus Steiniomyia Townsend runs here and has all
the essential characters of Hyleovus. It should be treated as a synonym of the
latter, but definite synonymy is not established at this time]

— Prosternum bare. Eyes bare or almost completely so. MM, spur vein or fold short,
not nearly as long as the section of M between m-—cu and the bend (Text-fig. 82).
One or both intermediate abdominal tergites without discal setae. Facial ridges
bare (except for the usual very few setulae immediately above the vibrissae)

HYSTRICOVORIA Townsend

[The Ethiopian and Oriental genus Afrovovia Curran (synonym Anavoria
Mesnil) runs here and has all the essential characters of Hystricovoria. It should
probably be treated as a synonym of the latter, but definite synonymy is not
established at this time]

Tribe THELAIRINI

This is a small but widely distributed tribe occurring in both Old and New Worlds.
The type-genus Thelaiva is found throughout the Palaearctic and Oriental Regions,
Africa and Madagascar, in New Guinea and in eastern Australia south to Tasmania,
and in the New World occurs southwards as far as Mexico; in the Pacific area the
genus is apparently unrepresented eastwards of New Guinea. The only genus found
so far in Australia, other than Thelaiva, is Halydaia, a very distinctive form occurring
in most of the Oriental Region eastwards to Japan, the Ryukyu Islands, New
Guinea, the Solomons and Australia (excluding Tasmania); at least one rare species
of Halydaia occurs also in western Europe. In the Old World Halydaia appears
to be rather disjunct from other thelairines, but the genus appears to be rather
closely related to the Neotropical genus Xanthodexia Wulp (syn. Minthodexia
Brauer & Bergenstamm) which is somewhat intermediate between Halydaia and
the more typical Thelairini.

The chief characteristics of the Thelairini are as follows. Eyes bare or virtually so (some
very minute hairs can sometimes be made out under high power examination); parafacials
bare; eyes extremely large and filling most of the sides of the head, gena correspondingly
very reduced and without a definite genal dilation (eyes coming so far down on the head
that the lowest point is usually below the level of the vibrissae); face and epistome flat, latter
invisible in profile; vibrissae usually level with epistomal margin (above the level in the
Oriental genus Prosheliomyia Brauer & Bergenstamm); antennae small or very small (length
less than half of eye-height) and antennal axis at or below level of eye middle; arista usually
short plumose or long plumose, sometimes pubescent; proboscis very short, mentum convex
on lower edge in profile, palpi well developed (a little flattened); prosternum, prosternal
membrane, and propleuron bare; two sternopleural setae (rarely three in 9); 3 + 3 dc setae;
pre-alar seta very small; two or three fost ia setae (rarely specimens may have only one);
infrasquamal hairs usually absent; scutellum with three pairs of very strong marginal setae
(basals, subapicals, and strong crossed horizontal apicals); pteropleural seta absent (except
in Prosheliomyia in which weak pteropleural always evident); fore coxa largely bare or fully
haired on inner anterior surface; mid tibia with a v seta; hind coxa bare posterodorsally;
hind tibia without a pv apical seta; upper surface of wing setulose at least on R,,; half way
to y-m, often veins FR, and Cu, extensively setulose (ft, setulose along all its length in both
genera occurring in Australia); cell R; open to the wing margin; bend of vein M moderately
sharp or rather evenly rounded, usually without M, appendix; second costal sector haired
ventrally; excavation of Ti + 2 variable, usually not quite reaching hind margin of tergite;

64 R. W. CROSSKEY

abdomen often with discal setae (discals present on T3, T4 and T5 in both Australian genera) ;
sternites concealed.

Attention should be called to the spelling of the generic name Halydaia. This
name in Tachinidae is usually spelled Halidaya, and this is reasonable from a
common-sense point of view since the genus was named in honour of the dipterist
Haliday. However, Egger (1856) was apparently under the impression that
Haliday’s name was spelt as ‘Halydai’ as he wrote ‘Ich habe sie zu Ehren des
englischen Dipterologen A. H. Halydai Halydaia genannt’, and nowhere in the
original description does the spelling Haliday or Halidaya appear. In this curious
circumstance it is open to question how Article 32 (a) of the International Code of
Zoological Nomenclature, 1961, applies, but Halydaia is held here to be a correct
original spelling; Egger was in the belief, it appears from the publication, that
Halydai was a correct spelling and the spelling of the generic name Halydaia was
therefore intentional (and cannot be held to be an inadvertent error).

It is necessary to have it established beyond doubt that Halydaza is the correct
spelling of the name of the Tachinid genus, because if not then the name Halidaya
in tachinids enters into homonymy with Halidaya Rondani in the Sepsidae, which
by unfortunate coincidence was published in the same year (1856) as Halydaia
Egger. (It is not evident at present whether Egger’s or Rondani’s was the earlier
of the two works, but as Halydaia Egger is held to be valid under the Code there is
no homonymy with Halidaya Rondani and it is irrelevent which name is the prior
one.)

KEy TO AUSTRALIAN GENERA OF THELARIINI

1 Antennae inserted at a level far below the eye middle (Text-fig. 36). Head in facial
view with the eyes converging ventrally so that the facial region is narrower than
the frons. Both sexes with 5 or 6 pairs of strong proclinate orbital setae
(Text-fig. 36). Arista pubescent. Eyes so large that they reach the peristome
and virtually obliterate the gena (Text-fig. 36). Scutellum without discal setae.
Interfrontal area reduced, much narrower than a parafrontal. Fore coxa haired
only on apical half of the inner anterior surface. Legs reddish yellow (except
sometimes for dark tarsi) : : ‘ : : : : HALYDAIA Egger
Antennae inserted at the level of the eye middle (Text-fig. 35). Head in facial
view with the eyes slightly or strongly diverging ventrally so that the facial region
is wider than the frons. @ without proclinate orbital setae, 2 with two pairs.
Arista short plumose. Eyes large but not reaching to the peristome, gena distinct
and about as wide as third antennal segment (Text-fig. 35). Scutellum with a
pair of semi-recumbent discal setae. Interfontal area wider than parafrontal.
Fore coxa haired on whole extent of the inner anterior surface. Legs black
THELAIRA Robineau-Desvoidy

Tribe MINTHOINI

This is an Old World tribe containing a few genera of very slender tachinids
with long narrow wings and elongate legs. A few New World forms have been
placed in the tribe but it is doubtful whether they correctly belong. A curious
feature of the group is the conspicuous flattening of the fore tarsi in the females;

TACHINIDAE OF AUSTRALIA 65

usually the female fore tarsi are flattened and dilated laterally, but in a few forms
(for instance the Australian Minthoxia) the flattening is from side to side only on
the basal part with the apical segments flattened and widened dorsoventrally.
The Australian fauna contains only two genera at present, Suwmpigaster and
Minthoxia; the first of these is a completely typical minthoine closely similar to
the type-genus, Mintho Robineau-Desvoidy (from the Palaearctic and Ethiopian
Regions), but the second has a facies rather different from typical minthoines and
may resemble them at all only convergently. In the original description of
Minthoxia it was placed in the Minthoini and compared with Tachinodexia Townsend
(an Oriental genus very similar to Swmpigaster) and it seems best to retain Minthoxia
in the Minthoini for the time being. Minthoxia is known only from Australia,
but Swmpigaster (originally described from Queensland) occurs widely in the
Oriental Region eastwards to New Caledonia and the Loyalty Islands. The genus
Megistogastropsis Townsend occurs in Papua, but has not yet been found in
Queensland; its occurrence there seems possible, however, and it is therefore included
in the key to the Australian genera of Minthoini.

The main characteristics of the Minthoini are as follows. Eyes usually bare or almost so;
parafacials bare; epistome flat; vibrissae about level with the epistomal margin; antennal
axis level with or below the eye middle; arista pubescent or plumose; proboscis short, palpi
fully developed; prosternum bare (except in Minthoxia), prosternal membrane bare; propleuron
bare; two or three sternopleural setae; acrostichal setae almost always reduced to 1 +0
or I + 1; 2 + 3 or 3 + 3 dc setae; pre-alar seta absent or minute; two or three fost ia setae,
pyrst ia seta usually absent; humeral callus almost always with two setae, occasionally a small
third seta in addition; infrasquamal hairs usually absent; pteropleural seta present, but fine;
scutellum rather flattened, without preapical (discal) setae, marginal setae not exceeding
three pairs (always without laterals); fore coxa largely bare on anterior surface or haired on
outer half of anterior surface; mid tibia with a v seta (often very small); hind coxa bare
posterodorsally, usually rather remote from abdominal base; hind tibia usually with distinct
pv apical seta; upper surface of wing with a few minute setulae on base of R,,;, veins otherwise
bare; cell R; open or short-petiolate; second costal sector bare or haired ventrally; bend of
vein M varied, usually rather abrupt, with or without /, appendix, sometimes gently rounded;
wings usually very long, apical part of Cw, shorter or much shorter than m-cu; Ti + 2 usually
not excavate to hind margin; some abdominal tergites usually with discal setae; sternites
(except basal sternite) concealed; abdomen usually very long and slender.

Key To AUSTRALIAN GENERA OF MINTHOINI

1 Eyes bare or virtually so. Prosternum bare. Acrostichal setae usually 1 + 0,
occasionally 1 + 1. Three post ia setae (sometimes weak, middle one undeveloped
in occasional specimen). Mid tibia with one ad seta. Very slender forms with
elongate legs ; : : ; ; : : : : : ; :

— Eyes conspicuously haired. Prosternum setulose. Acrostichal setae variable but
numerous, at least 2 + 3. Two post ia setae (both strong, wide apart). Mid
tibia with two ad setae. Not such slender forms, legs (especially fore tarsi)
robust . : t : : : ‘ - : : MINTHOXIA Mesnil

2 Abdomen with strong discal setae on last visible tergite (T5), similar to those on
T3 and T4. Arista plumose (Text-fig. 32). Second costal sector bare ventrally.
Scutellum with two pairs of marginal setae (either basals or apicals or both
present in addition to subapicals). Bend of vein M sharp, with or without M,
appendix. Pre-alar seta almost always completely absent SUMPIGASTER Macquart

E

66 R. W. CROSSKEY

— Abdomen without discal setae on T5. Arista pubescent. Second costal sector
haired ventrally. Scutellum with only one pair of marginal setae (the subapicals),
basal and apical setae absent. Bend of vein M forming an evenly rounded curve
without trace of appendix. Pre-alar seta almost always present, though small.
[Indonesia & New Guinea] : : : . MEGISTOGASTROPSIS Townsend

Tribe NEMORAEINI

This small tribe is confined to the Old World and contains only the genera
Nemoraea Robineau-Desvoidy (of which Pvotonemoraea Baranov and Dexiomima
Brauer & Bergenstamm are now treated as synonyms) and Chaetolydella Villeneuve.
The latter genus is African and is not really distinguishable from Nemoraea, as
van Emden (1960) states, but this is not the place in which to establish synonymy.
Nemoraea is widely distributed throughout the Palaearctic Region (including
Japan), Africa, and the Oriental Region, but until now has not been reported from
Australia. One undescribed species is now known from New South Wales and
Queensland; this species has a mainly orange-red abdomen and a black-and-yellow
striped scutum and closely resembles N. ornata (Bigot) from the Oriental Region
(it also has a superficial appearance very reminiscent of the New Zealand genus
Protohystricia Malloch, which belongs in the local New Zealand tribe Occisorini).
In New Guinea the Nemoraea fauna contains at least four undescribed species, and
there are some large and spectacular species still undescribed in the Oriental fauna.

The numerous species of Nemorvaea tend to fall into two moderately distinct
categories, one containing species in which the whole surface of the lower calypter
is haired and the second costal sector bare ventrally and the other containing
species in which hairing of the upper surface of the lower calypter is confined to the
outer edge and the second costal sector is haired below (the Australian species falls
into the latter category). The name Nemoraea strictly applies to the first group,
and the names Dexiomima and Pyrotonemoraea to the latter group. Many
intermediates exist, however, and the current treatment of all the forms as comprising
the single genus Nemoraea is appropriate; examples of intermediate species, falling
between the two main categories, are N. echinata Mesnil from Burma in which the
outer half of the lower calypter is haired, and N. dotata (Walker) from Celebes in
which the lower calypter has hair only on the outer margin but in which the second
costal sector is bare on its ventral surface.

The chief characters of the Nemoraeini are as follows. Eyes haired; parafacials bare;
arista micropubescent; face usually with a distinctly raised central ridge which is rounded
on its anterior surface (not forming a sharp carina); genal dilation bearing unusually strong
setulae or even setae as strong as the peristomal setae (Text-fig. 38); epistome flat, invisible
in profile; vibrissae level with or only very little above the level of the epistome; inner vertical
setae crossed (sometimes subparallel in 3); outer vertical setae unusually weak in Q, often
not differentiated from postocular row; palpi unusually dorsoventrally flattened, band-like;
prosternum, prosternal membrane and propleuron bare; three post ia setae; 3 + 3 or 3 + 4 de
setae; two or three sternopleural setae; infrasquamal hairs present or absent; pteropleural
seta present; fore coxa completely haired, or almost so, on its inner anterior surface; mid
tibia with a v seta; hind coxa bare posterodorsally; hind tibia with a small (rather

TACHINIDAE OF AUSTRALIA 67

inconspicuous) pu seta; wing with only a few setulae basally on R,,;, veins otherwise bare;
cell R; open to the wing margin; bend of vein M very sharp and with a conspicuous M, spur
vein or fold; second costal sector bare or haired below; abdominal Tr + 2 excavate to its
hind margin; ¢ hypopygium usually with some rather long strong setae visible in situ;
sternites mainly or partly overlapped by tergites, but at least exposed at their apices.

Tribe LESKIINI

This tribe is in much need of a reclassification of the genera on a world basis, as
the faunae of the Old World and New World regions contain forms that are virtually
indistinguishable yet have not been associated in the same genera. The rich
Neotropical fauna has been subject to excessive generic splitting, which has badly
obscured the relationships existing within this local fauna and has made it difficult
to relate the South American leskiines with Old World forms. This is unfortunate,
as even a casual comparison shows that species occur in the Pacific islands and
Australia, for example, that appear to be congeneric with New World forms. Bezzi
(1928) placed a Fijian species in Sipholeskia, and Curran’s Australian species
Demoticus certima is here placed in the same genus after comparison with the
type-species of Sepholeskia. Another of Bezzi’s (1928) Fijian species, ‘Rhinomyiobia’
minuta, is so closely similar to the Neotropical species Mvyobiopsis diadema
(Wiedemann) that it ought perhaps to be assigned to Myobiopsis Townsend or
one of the related South American genera, and the same comment applies to a small
unidentified leskiine from Queensland that is exceedingly similar to minuta Bezzi
(this species is run out in the key that follows as representing an unidentified genus).

In Australia and Tasmania the Leskiini are rather well represented, though at
present most of the forms are undescribed and for several of them it is impossible
at present to decide upon a suitable generic assignment. Six named and identifiable
genera occur in the area, of which two are non-endemic (Demoticoides and
Sipholeskia). The others — Apatemyia, Toxocnemis, Exechopalpus and Rhinomyobia
— are endemic genera not found outside Australia and Tasmania (a careful comparison
with Leskiini from other regions has shown that only Rhinomyobia has characters
fitting closely with any extra-limital forms). Bezzi (1928) placed two Fijian
species in Rhinomyobia, but Townsend (1939 : 300) is correct in stating that neither
of them truly belongs in this genus; it may here be noted that Townsend (loc. cit.)
is also right that Rhinomyobia transversalis Malloch is wrongly assigned to the
genus (this species, from Queensland, is undoubtedly congeneric with ‘Rhinomyiobia’
plumifera Bezzi from Fiji but there appears to be no available generic name to
apply to these species at present).

It is difficult to define the Leskiini with much precision, especially as the tribe
contains a few species in which the prosternum is haired, or the propleuron is haired,
and seemingly closely allied forms may or may not possess a pv apical seta on the
hind tibia, and differ in the arrangement of the scutellar bristling. Typically,
however, the tribe includes rather slender long-legged forms in which the epistome
is projecting, the arista long-pubescent or plumose, the eyes bare, the fost dc setae
number three, the mediotergite lacks infrasquamal hairs, the scutellum has only two

68 R. W. CROSSKEY

pairs of strong setae, the bend of vein M lacks an appendage, cell R; is open to
the wing margin or only just closed, and there are two or three stf/ setae. Some
forms occur, less typical of their tribe, that have hairy eyes, four post dc setae,
infrasquamal hairs, and three pairs of scutellar setae. Pallid coloration and reddish
yellow legs are rather commonly found in the group, and there is a tendency for
the tibiae of the males of the really long-legged forms to become extremely slender
and sinuous with associated reduction of the bristling (here it should be noted that
the mid tibia in Leskiini appears always to possess a submedian v seta but that this
is often extemely small in long-legged males).

The endemic genus Exechopalpus possesses extraordinarily long palpi, similar to
those of the Neotropical genus Spathipalpus Rondani, and the proboscis itself is
extremely slender. Throughout the Leskini there is a marked tendency for the
proboscis to be unusually slender, elongate, almost stylet-like, with the labellae
correspondingly reduced, and even in forms in which the proboscis is short it is
usually found that the mentum is parallel-sided in profile rather than convex
on its lower edge.

The genus Bezziomyiobia Baranov from the Solomon Islands has not been
found in Australia, but is included in the following key to genera to show its main
features. It is extremely close to Demoticoides Mesnil, and the latter ought perhaps
to be sunk as a synonym of Bezziomyiobia (it is beyond the scope of the present work
to consider this further).

KerEy To AUSTRALIAN GENERA OF LESKIINI

1 Scutellum with only two pairs of strong marginal setae, the basals and subapicals;
apical scutellar setae minute and hair-like or absent, if weakly developed then
directed half-upwards. Subapical scutellar setae not inserted very widely apart,
distance between their bases subequal to or less than that between a subapical
seta and its corresponding basal seta : 2

— Scutellum with three pairs of strong marginal = a oak of strong horizaatel
crossed apical setae present in addition to the basals and subapicals. Subapical
scutellar setae inserted very widely apart, distance between their bases about twice
as great as that between a subapical seta and its corresponding basal seta (or even
more than this) . 8

2 Palpi exceptionally Sunes length sleet eaeal to that of ead at level a the
epistome; palpi projecting beyond epistome by a distance about equal to the
antennal length (when proboscis retracted). Fore tibia with some strong ad
setae. Mid tibia with three or four strong ad setae and with three strong p
setae. Hind tibia with a pu apical seta . : . EXECHOPALPUS Macquart

— Palpi normal, length conspicuously less than that of head at level of the epistome;
palpi not or only a little projecting beyond epistome (when proboscis retracted)
(Text-fig. 41). Fore tibia without ad setae (a few inconspicuous ad setulae on
basal half in Rhinomyobia). Mid tibia with one or two ad setae and nearly
always with two p setae (some Towxocnemis-like forms with a very small third p
seta basad of the normal two). Hind tibia without a pu apical seta (except in
Sipholeskia) . : : : : : : 3

3 Abdomen with Ti + 2 ae to Be pad eee Second costal sector bare
ventrally. Humeral callus with three or more setae in line (sometimes not
clearly differentiated from long hair). More robust forms with subovate or
subconical abdomen } : ; : E : : : 3 : : 4

“I

TACHINIDAE OF AUSTRALIA 69

Abdomen with Tr + 2 excavate only on its basal half. Second costal sector haired
ventrally. Humeral callus usually with only two strong setae (sometimes a weak
third seta set forwards of main two or mesad of inner one of main _ Slender
forms with rather long narrow abdomen . 4

Pteropleuron with a strong pteropleural seta in addition to the hairing. Parafacials
bare or at most haired only on upper half. Eyes bare or almost so. Infrasquamal
hairs absent. Arista bushy plumose or micropubescent . 5

Pteropleuron without any seta in addition to the hairing. Paratncials fully fenced.

Eyes haired (hairing very short and sparse in some specimens). Infrasquamal
hairs present [careful examination needed]. Arista bushy plumose
APATEMYIA Macquart

Propleuron bare. Proboscis short, mentum shorter than eye-height and wider in
profile than third antennal segment. Pleural eget of thorax with black hair.

Hind tibia of both sexes with two pd setae : 6

Propleuron haired. Proboscis very slender, calories longer fhan a height aud
narrower in profile than third atennal segment. Pleural regions of thorax with
yellowish white hair. Hind tibia of f with one pd seta (unknown for Q, possibly
two pd in this sex) : . TOXOCNEMIS Macquart

Parafacials totally bare. Arista with short bushy plumosity or long pubescence.

Rows of frontal setae irregular at lower ends where frontal setae form a rather

dense bunch on the profrons (especially in g): 4 abdomen with long fine erect or

semi-erect hairing. Intermediate abdominal tergites often with erect (rather

variable) discal setae. : . Undetermined genus
[Running out here is a complex of forms with exactly similar appearance in

most respects to Toxocnemis (head profile as Text-fig. 43) but differing by the

characters shown in the key, most notably by having the propleuron bare. This

complex will either require a new genus or will need to be placed in Toxvocnemis on

a redefined basis]

Parafacials haired on upper half or at least with a few hairs immediately below the
lowest frontal seta. Arista micropubescent (appearing almost bare at first glance).
Frontal setae in one regular row on each side, not duplicated into a bunch of setae
on the profrons. dg abdomen with short recumbent hair. Intermediate abdominal
tergites without discal setae . ; . RHINOMYOBIA Brauer & Bergenstamm

Wing vein R,,; completely bare. Proboscis short, mentum much shorter than
eye-height. Hind tibia without pu apical seta. Vibrissae level with epistomal
margin. Fore tibia without pd setae. Hind tibia with one pd seta Undetermined genus

[Running out here is an unidentified species seen from A.C.T. that cannot be
satisfactorily assigned to a genus at present. The species is congeneric with
‘Rhinomyiobia’ minuta Bezzi from Fiji.]

Wing vein R,,, with several setulae on basal node and usually for some distance
along the vein towards y-m. Proboscis long and very slender, mentum longer
than height of the head. Hind tibia with a small pv apical seta. Vibrissae
distinctly above level of epistomal margin. Fore tibia with some small pd setae
or setulae distinct from the hairing. Hind tibia with two or more small pd
Setae: « : : : é : : , ‘ . SIPHOLESKIA Townsend

Prosternum bare. Arista micropubescent. Eyes haired (hairing very short and
inconspicuous in 9). Abdominal T5 with a transverse discal row of very strong
setae in addition to the apical row. Humeral callus with the three main setae
distinctly arranged inatriangle . : : . 9

Prosternum setulose. Arista with long ilerettosiey. ‘Byes. bare. Abdominal T5
without a discal row of setae, only with apical setae. Humeral callus with the
three main setae standing almost in a straight line . : . New genus

[Running out here are Rhinomyobia transversalis Malloch from Queensland and
R. plumifera Bezzi from Fiji, together with some unnamed forms from New

70 R. W. CROSSKEY

Guinea, none of which correctly belongs in the genus Rhinomyobia. There appears
to be no described genus to which these species can be correctly assigned, and a
new genus will probably be needed for their appropriate placement. ]
9 Mid tibia with two ad setae. Legs and palpi dark brown. [Solomon Islands]
BEZZIOMYIOBIA Baranoyv
~— Mid tibia with one ad seta. Legs (except tarsi) and palpi reddish yellow. [India &
Queensland] ; : ‘ : ; ‘ : . DEMOTICOIDES Mesnil
[This genus is so similar to Bezziomyiobia that the name ought perhaps to be
treated as a synonym. ]

Tribe ERNESTIINI

Four endemic Australian genera are here placed in this tribe, but it is doubtful
whether the placements of two of them, Neximyia and Amphitropesa, will be upheld
by further work. The affinities of these two genera are very obscure, and the fact
that they are so far known only from very few female specimens makes it specially
difficult to determine their true relationships; Neximyia at least appears to be
rightly placed in the Tachininae, and probably does truly belong somewhere near
the ernestiine-linnaemyine complex, but Amphitropesa is much more problematical
and may not really belong in the Tachininae at all (its total facies, much reduced
dorsal thoracic bristling, and stongly clubbed female palpi give it some resemblance
to certain Proseninae).

The placement of Chlorotachina in Ernestiini appears to be almost certainly correct.
In all its characteristics it conforms to the Ernestiini as the tribe is recognized
in the Eurasian fauna, and the dark metallic blue-black or metallic greenish blue
appearance of most forms conforms with that of similar metallic forms in the
Palaearctic Region. The genus Macrochloria, although not immediately obvious
as an ernestiine, is best placed in the Ernestiini as its characters closely accord
with those of Chlorotachina.

The main characteristics by which Chlovotachina and Macrochloria concur with Eurasian
Ernestiini are as follows: epistome very prominent, vibrissae inserted well above epistomal
margin; parafacials bare; palpi moderately well developed; eyes densely long-haired; three
postsutural za setae (cf. Parerigonini); scutellum with four pairs of very strong marginal setae;
mid tibia with more than one ad seta and both sexes with a v seta (¢ lacking a v seta in typical
Parerigonini); hind coxa bare posterodorsally; wing with bend of vein M very sharp and
provided with at least a trace of an 17, stump or spur vein.

The genera Neximyia and Amphitropesa, here placed as a temporary measure in
this tribe, do not conform with all of these characteristics, as will be evident from
the following key to genera.

KEY TO AUSTRALIAN GENERA OF ERNESTIINI

1 Eyes densely haired. Presutural acrostichal setae well developed. Prosternum
haired (except in occasional specimen of Macrochloria). Wings clear hyaline : 2

— Eyes bare (some very sparse short macroscopic hairs may be visible under high
power). Presutural acrostichal setae absent or hair-like. Prosternum bare.
Wings with the membrane partly brown and partly yellow : : : : 3

TACHINIDAE OF AUSTRALIA 71

2 Upper half of face raised medially into a definite ridge-like or knob-like keel beneath

the antennae, the keel easily visible when head seen in profile. Abdomen with

dense white pollinosity on each side of the last visible tergite (T5), this tergite

therefore contrasting in appearance with shining remainder of abdomen. Thoracic
dorsum not shining . : : MACROCHLORIA Malloch

— Upper half of face slightly swollen medially but not raised into a keel, face therefore

almost completely invisible in profile. Abdomen entirely shining, if some

whitish pollinosity present then rather evenly distributed and very inconspicuous.
Thoracic dorsum distinctly shining . . CHLOROTACHINA Townsend

3 Parafacials bare. Propleuron bare. Wing mth cell R; open at the wing margin.

Form with remarkable pattern, as follows: dorsum ee thorax very pale greyish

yellow with six large strongly contrasting black marks (sublateral pair on prescutum

and a lateral and a sublateral pair on scutum); abdominal dorsum black on

Tr + 2 and T3 (except for pair of pale greyish yellow marks anteriorly on outer

edges of T3) and densely covered with pale greyish yellow pollinosity on T4 and

T5, T4 with three bold black spots (median spot and a pair of lateral spots that

continue round to ventral surface). Apical half of wing irregularly smoky

brownish, the brown colour nowhere sharply contrasting with hyaline areas

AMPHITROPESA Townsend

— Parafacials haired (Text-fig. 45). Propleuron haired. Wing with cell R; closed at or

a little before the margin. Without such pattern, but marked as follows: dorsum

of thorax with shining black prescutum and scutum contrasting with yellow or

orange scutellum, humeral calli and notopleura, the pale areas with pale golden

pollinosity; abdomen bright orange with a broad black fascia across the middle

covering T4 and apical part of T3, usually also three small black spots on T5 and

median depression of Tr + 2 dark. Wing with a sharply defined but irregular

brown cross-band, and with the membrane clear hyaline distally to the cross-band
NEXIMYIA Crosskey

Tribe PARERIGONINI

This rare and aberrant group, first defined by Mesnil (1966 : 888), occurs in
Eurasia, New Guinea and Australia. Its relationships with other tribes are at present
very obscure, but the unusual modification of the male postabdomen, the unusual
male genitalia, and the variously modified female terminalia, should shed some
light on the true affinities when they are studied in detail. Superficially the New
Guinea forms much resemble some Goniinae, and Australian forms like Zita have
a general appearance, head facies and scutellar bristling much like some Proseninae;
there is also some superficial resemblance to the Linnaemyini, mainly because of
the prominent epistome and high vibrissae, and Pygidimyia has the colour and
form much as in Chaetophthalmus. The Australian genera are known only from
very few specimens, but these include representatives of some new species.

The main characteristics shown by the Australian Parerigonini are as follows. Epistome
prominent and vibrissae inserted high above the epistomal margin; parafacials bare; genal
dilation well developed; g without proclinate orbital setae, 2 with a pair of outwardly directed
prevertical setae; inner vertical setae often convergent or crossing; upper occiput without
black setulae behind the postocular row; prosternum bare; propleuron usually haired; humeral
callus with three strong setae in a triangle, and one smaller seta set forwards of the inner two
of the triangle; 3 (4) + 3 dc setae; 0 + 2 ia setae (the two post ia setae very strong and
normally inserted in a position that suggests that a third post ia is never developed); three

72 Ke) We ‘GROSSE,

or four stp] setae (2 + 1 or 3 + 1); infrasquamal hairs present; pteropleural seta present or
absent; mid tibia without a v seta in the g, with a v seta in the 9 [this sexual difference
apparently constant to judge from limited material seen]; hind coxa bare posterodorsally;
hind tibia without pd preapical seta and without pv apical seta; wing with only a few weak
hairs on basal node of R,,;, veins otherwise bare; cell R; open to the wing margin; bend of
M rather abruptly angulate but without M, spur or fold; abdomen of both sexes unusual,
in ¢ To and T7 + 8 exposed and forming a conspicuous vertical declivity at the end of the
abdomen, and in 9 postabdomen strongly developed and recurved with elaborately modified
Ovipositor (sometimes resembling that of some Phasiinae such as Cylindromyiini); abdominal
sternites exposed.

The strange genus Awustralotachina Curran does not completely conform with the
characters cited above, as it has sparse black hairs on the upper half of the occiput,
has four post dc setae, the ¢ has a submedian v seta on the mid tibia, there is an
extra pair of scutellar setae, and the bend of vein M is rather gently rounded, but
it seems nevertheless best to place it with the parerigonines until its affinities can
be more reliably determined; its head profile is shown in Text-fig. 42.

KEY TO AUSTRALIAN GENERA OF PARERIGONINI

1 Scutellum with three pairs of very strong setae (basals, subapicals and crossed

apicals). o + 2zasetae. Propleuron usually haired. 3 + 3 dcsetae sili =
weak fourth prst dc seta) : 2

— Scutellum with four pairs of strong setae G sa ue enormous laterals presente in

addition to basals, subapicals and crossed apicals). o + 3 ta setae (a weak

hair-like post ia seta present in front of the main two). Propleuron bare. 3 + 4
dc setae : : : 5 : . AUSTRALOTACHINA Curran

Abdomen with Tr 3 and T3 normal, ‘GE fused, with excavation of Tr + 2

reaching the hind margin. Barette haired on anterior half. Posterior thoracic

spiracle not exceptionally large, about as long as the barette or only a little more.
Both thoracic spiracles brown or blackish. Legs and all thoracic hair black 5 3

— Abdomen with Tr + 2 and T3 fused into a single very large composite tergite in

which the excavation is confined to the basal part. Barette bare. Posterior

thoracic spiracle exceptionally large, very much longer than the barette. Both

thoracic spiracles pale yellowish. Legs reddish yellow (except for apical darkening

of the tarsi) and hair of sides of the thorax golden orange or yellow

PYGIDIMYIA Crosskey

3 Eyesdensely and conspicuously hairy. Epistome usually only moderately prominent,

vibrassae inserted above the epistomal margin by a distance not exceeding the
length of the second antennalsegment . ; : : LEVERELLA Baranov

— Eyes bare or almost so (at most only with very sparse inconspicuous and microscopic

hairs). Epistome strongly projecting, subnasute, vibrissae inserted above the

epistomal margin by a distance exceeding the length of the second antennal
segment (Text-fig. 46) . : : : . : ; : : ZITA Curran

Nn

Tribe LINNAEMYINI

The Linnaemyini include forms that are superficially very like the Tachinini
but differ by having the dorsal surface of the hind coxa bare. All the Australian
forms have the eyes densely haired, and this helps to separate them at a glance

TACHINIDAE OF AUSTRALIA 73

from the Australian tachinines in which the eyes are always bare. Separate tribal
status for Linnaemyini and Tachinini is in reality scarcely justified, but the Australian
forms at least can be readily differentiated and separate tribal status is recognized
for purposes of the present work. The following principal features are shared by
the two groups: Epistome prominent or subnasute; vibrissae inserted well above
level of epistomal margin; scutellum with similar bristling which usually includes
erect subspiniform preapical setae; mid tibia with several ad setae and a submedian
v seta; hind tibia with a pv apical seta; cell R; open; bend of vein M sharp and
provided with an M, extension either as a definite spur vein or at least as a wing
fold; pteropleural seta strong; abdomen with Tr + 2 excavate to its hind margin
and with the sternites largely exposed; upper occiput without black setulae behind
the postocular row.

Three genera are known from Australia, at least one of which (Chaetophthalmus)
occurs also in Tasmania. This genus up to now was believed to be entirely
Australian, but undescribed species are now available from New Guinea and New
Caledonia. Chaetophthalmus is the dominant linnaemyine genus in Australia, and
the other genera appear to be uncommon to judge from material so far known;
one of these is the widespread Old World and North American genus Linnaemya,
and the other is the monotypic genus Apalpus which is known only from South
Australia and Western Australia. The three genera can only be reliably separated
by the vestiture, or lack of vestiture, on the parafacials, and there is some
doubt whether generic status is fully justified. It may be noted that they all have
3+ 3dc setae (Text-fig. 61) and in this respect differ from the tachinines in
Australia which all have 4 + 4 dc setae or more; the propleuron is always bare,
and this character (as well as the hairy eyes) is useful in differentiating Chaetoph-
thalmus from the tachinine genus Microtropesa, which has some _ superficial
resemblance to it. All the Australian Linnaemyini have very reduced palpi.

KrEy TO AUSTRALIAN GENERA OF LINNAEMYINI

I Parafacials haired. Normally two stpl setae. o + 3 ia setae [a prst ia seta probably
sometimes occurs but has not been seen]. Epistome extremely prominent and
nasute. Vibrissae inserted very high above the epistomal margin, distance from
insertions to level of the epistomal margin as great as or greater than the length
of the second antennal segment. Prosternal membrane and anterior part of
prosternum usually finely haired (hairs if present may be pale and very
inconspicuous). Abdomen rather rotund, appearing noticeably wider than
thorax to naked eye : c : : : c ; : : -

— Parafacials bare. Normally three stp/ setae. 1 + 3 ia setae [prst ia seta probably
sometimes absent]. Epistome prominent but not strikingly nasute. Vibrissae
inserted above the level of the epistomal margin by a distance less than the length
of the second antennal segment. Prosternal membrane and prosternum bare.
Abdomen more elongate, not noticeably wider than thorax to naked eye

LINNAEMYA Robineau-Desvoidy

2 Parafacials without any strong setae in addition to the hairing (Text-fig. 44)

CHAETOPHTHALMUS Brauer & Bergenstamm

— Parafacials each with one strong incurved seta near the lower end in addition to the

hairing (see fig. 17 in Malloch, 1929b : 316) : d : : APALPUS Malloch

Ne

74 R. W: CROSSKEY

Tribe TACHININI

In Australia the Tachinini are represented mainly by the endemic genus
Mucrotropesa, which is readily distinguished from other tachinine (except Paratropeza
Paramonov) genera by the possession of long dense hair on the pleurotergite (the
supraspiracular convexity just dorsal to the hind thoracic spiracle). This genus
includes several distinctive species with patterned abdomen and smoky brown
and yellow wings, but also some species of more humdrum appearance. The only
two other genera known in Australia are non-endemic, Cuphocera occurring very
widely throughout the Old World and Ervistaliomyia in the south-east Asian
archipelago, Malaya and New Guinea (the latter genus is probably confined in
Australia to the Northern Territory and Queensland); Evistaliomyia, it should be
noted, is very closely allied to the genus Servillia Robineau-Desvoidy (which is
abundantly represented throughout the Oriental Region) and perhaps ought not
to be recognized as a valid genus. In general, the tachinine fauna of Australia is
rather impoverished, in contrast to that of the Palaearctic and Oriental Regions
where it is so richly developed.

The genera represented in Australia have the following features in common that aid in
tribal recognition. Parafacials covered with long fine hair; second antennal segment unusually
elongate (and the third not or hardly longer than the second); vibrissae inserted far above
epistomal margin; eyes bare; epistome very strongly projecting; prosternum bare; 4 + 4 de
setae (Text-fig. 62) (sometimes with additional dc); scutellum often with erect spiniform setae
or setulae just before the apex; no infrasquamal hairs; mid tibia with many ad setae and with
a v seta; hind tibia with a strong pu apical seta; hind coxa haired on the posterodorsal surface;
basal node of R,,; with only a few weak hairs, veins otherwise bare; second costal sector bare
below; cell R,; open; strong pteropleural seta (often doubled); abdomen with T1 + 2 excavate
to hind margin; abdominal sternites exposed and bearing stiff, often spiniform, setae; bend
of vein M abrupt and with a weak fold extending at least slightly from the bend towards the
wing margin.

KEY TO AUSTRALIAN GENERA OF TACHININI

1 Parafacials without strong setae in addition to the hair. Propleuron haired.
I+272asetae. Palpi well developed (though sometimes very acy 3 without
proclinate orbital setae . ‘ 2

— Parafacial armed with two or three strong setae (standing on lower end near the ea
I + 3 72asetae. Palpi vestigial. @ with or without proclinate orbital setae

CUPHOCERA Macquart

2 Convexity above posterior thoracic spiracle (pleurotergite) bare. Three strong
supra-alar setae. One sternopleural seta [o + 1 stpl in specimens seen, possibly not
constant]. Parafrontals shining black. Forms with bee-like facies and unusually
short wings . ; 2 : ERISTALIOMYIA Townsend

— Convexity above posterior thoracic spizacle (pleurotergite) with dense vestiture of
long hair. Two or three sternopleural setae. Parafrontals yellow to dark reddish
orange. Forms without such facies and with wings not unusually short in relation
to body size . ‘ 3 . : ‘ ‘ , MICROTROPESA Macquatt

SUBFAMILY GONIINAE WITH KEYS TO THE TRIBES AND GENERA

This enormous subfamily includes the forms commonly thought of as the ‘higher
Tachinidae’, and many of them have the pre-alar seta very large; this character

TACHINIDAE OF AUSTRALIA 75

appears clearly to be a derived (apomorphic) one, and is not found to the same
degree elsewhere in the Tachinidae. Some authors have in the past recognized
two subfamilies within the group, the Exoristinae (= Tachininae sensu van Emden)
with a small pre-alar seta, and the Goniinae with a large pre-alar seta, but specialists
now seem agreed that it is only justified to recognize all as a single subfamily
(Goniinae); the group as a whole is rather uniform in the genital structure of the
males and intermediate forms exist between the more typical elements of the
exoristines and the goniines on external characters. The external facies within the
Goniinae is extremely diverse, though not to the extent shown by the Tachininae,
and it is not easy to define the subfamily (especially as it is not unified by parasitizing
only one order of insects, the hosts including forms as diverse as adult beetles,
larval Lepidoptera and grubs of paper-making wasps). Nevertheless the following
characteristics of the external adult morphology are typical of the subfamily and
help towards defining it.

Head without a facial carina; head of g never holoptic and uppermost eye facets not
enlarged; rows of frontal setae descending usually to about on a level with the middle of
the second antennal segment; reclinate orbital setae nearly always present, 2 rarely with
outwardly directed prevertical setae; inner vertical setae normally subparallel (very rarely
crossing); vibrissae well developed; arista normally micropubescent, rarely plumose; palpi
almost always fully developed (papilliform in some species of Stomatomyia, Spoggosia, etc.) ;
thoracic and abdominal chaetotaxy usually strongly developed; prosternum haired or setulose
(bare in a few forms, especially in Blondeliini); prosternal membrane bare; humeral callus
with two or more setae; at least 2 + 2 dc setae, usually more; normally three post ia setae
(two only in a few forms or aberrant specimens); pre-alar seta present, small or large; two or
more sa setae, though hind one sometimes weak; postalar callus with two setae; scutellum
with at least two pairs of marginal setae; 2—4 (5) stpl setae; infrasquamal hairs almost always
absent (present at least in a few Blondeliini); fore coxa bare on much of its inner anterior
surface; mid tibia with or without a submedian v seta; hind coxa almost always all bare
posterodorsally, but a few forms (such as many Carcelia spp.) with one or two fine setulae;
hind tibia almost always without pu apical seta, usually with two dorsal preapical setae but
sometimes with a pd preapical seta in addition; abdomen with Tr + 2 excavate to its hind
margin (a few exceptions including all Siphonini) ; sternites of the abdomen normally concealed
or mainly so; hair of lower half of the occiput and the postbuccae always pale.

Some of the tribes here accepted are very weakly defined and probably ought
not to be recognized as valid, but it is best to treat them as valid until a better
classification can emerge from a large-scale study on a world basis using new or
improved criteria. This comment applies particularly to the Carceliini, Sturmiini,
Winthemiini and Eryciini, which merge rather imperceptibly into one another
and are only separable on rather unconvincing grounds.

Herting (personal communication) considers that the multifarious genera of the
Goniini-Carceliini-Sturmiini-Eryciini complex should be aggregated into two tribes
(for which the names Eryciini and Goniini would be nomenclaturally correct)
according to whether they have an ovolarviparous or a microoviparous reproductive
habit. Such a course has much to commend it insofar as it would probably
reflect the real phylogeny more accurately than the present tribal system. But
it is impossible to adopt such a system as yet for the Australian fauna, in which
the reproductive habit of most of the genera remains unstudied. Furthermore,

76 R. W. CROSSKEY

a separation into two tribes on the basis of the reproductive habit presents practical
difficulties of identification (since the external adult morphology of the genera
included in the redefined Goniini and Eryciini would not allow clear diagnosis,
nor would it lend itself to tribal key construction).

KeEy To AUSTRALIAN TRIBES OF GONIINAE

1 Pre-alar seta long and strong, longer than the first post 7a seta and usually longer

than the first post dc seta (except in Bactromyiella). Sternopleuron often with

four stpl setae. Second costal sector bare ventrally. eat scutellar setae

never converging to enclose the apicals . - : 2
— Pre-alar seta small and weak, shorter than ie ae as 1a fa (except in

Phorocerosoma) and normally much shorter than the first post dc seta. Second

costal sector bare or haired ventrally. Sternopleuron with fewer than four stp/

setae. Subapical scutellar setae sometimes convergent and enclosing the

apicals : : : : - - : : : : : : 7
2 Ocellar setae ses Ses, Scutellum with a pair of stiff erect divergent preapical

setae. rons very wide in both sexes, g with outer vertical setae like the 9.

Eyes bare. 3 + 4 dc setae ; : ; : ; : GONIINI (p. 96)
— Ocellar setae proclinate or absent (slightly divaricate in Pavagonia). Scutellum

usually without stiff erect preapical setae just before the tip. Frons usually

not exceptionally wide, g rarely with distinct outer vertical setae. Eyes bare

or haired. Various complements of dc setae . : 3
3. Eyes relatively very large so that the gena is reduced to a narrow an below the
eye that is not as wide as the profrons (Text-figs 48, 52 & 53) . 4

— Eyes occupying relatively less of the side of the head, gena usually at least slightly
wider than the profrons (when doubtful check Winthemiini), often conspicuously

so (Text-figs 50 & 51) . : : é ; : : : 4 : : 5
4 Propleural seta present. Legs blackish brown or mainly so, basicosta usually
dark (yellowish in a few forms). [Parasites of Lepidoptera} . CARCELIINI (p. 89)

— Propleural seta absent (except in Kovalliomyia). Legs uniformly pale orange-yellow
or at most only femora brown, basicosta clear yellow-orange. [Parasites of
vespoid wasps] . ‘ . ANACAMPTOMYIINI (p. 90)
5 Barette completely fe Hagnecal callus with five strong setae (at least in the
6) of which the three main ones stand in a triangle. Eyes densely haired.
Hind tibia of g with a long close-set ad fringe. g without definite reclinate
orbital setae. Scutellum with strong horizontal crossed apical setae and with the
subapical setae inserted far apart (distance between their bases at least as great
as that between a subapical seta and its corresponding basal seta) (Text-fig. 72)
WINTHEMIINI (p. 88)
— Barette only haired at its anterior end (some exceptions: fully haired in Bactromyiella)
or entirely bare. Humeral callus with fewer than five setae, normally three setae
in a straight line and a fourth seta set forwards of the basal line of three
(sometimes only three setae in line or ina triangle). Eyes bare or haired. Hind
tibia with or without such fringe in the ¢. Arrangement of scutellar setae varied,
as above only in a few Sturmiini . 6
6 Vibrissae inserted at a level distinctly above the ppeeaeel eee inate ouilg
slightly above). Always 3 + 4 dc setae. Inner posterior angle of the lower
calypter well developed, and inner margin of lower calypter abutted closely
against the scutellum. 4 often with well developed close-set ad fringe on hind
tibia and sometimes with dense secondary sexual hair fascicles on venter of
abdominal T4 . : : : : : : ; . STURMIINI (p. 91)

Io

TACHINIDAE OF AUSTRALIA

Vibrissae usually inserted about on a level with the epistomal margin (if above then
other character not fully fitting or doubtfully so). Dorsocentral setae sometimes
3 + 4 but other complements occurring. Inner margin of lower calypter usually
not closely following edge of scutellum, inner posterior angle of the calypter
usually rather rounded. ¢ without a oa, close-set hind tibial fringe and

without abdominal hair fascicles. : : ERYCIINI (p. 96)

Scutellum with three pairs of very strong setae in which the apical setae are crossed
and horizontal and the subapical setae are very wide apart and inserted well
forwards on the sides of the scutellum (Text-fig. 68); no lateral scutellar setae.
Basal node of R,,; with one strong setula. Parafrontals in both sexes with a
long series of strong proclinate orbital setae. Antenna with the anterior tip of
the third segment forming a sharp point. Bend of vein M widely obtuse, M,
unusually straight and cell Rk; normally closed at the wing marginor short-petiolate
(Text-fig. 88). Second costal sector haired ventrally. 2+ 3 dc setae.

[Parasites of Orthoptera Acridoidea] . : : ACEMYINI (p. 78)

Scutellum almost always with different conformation of setae, lateral setae usually
present; if scutellar bristling similar then other characters not fitting. Basal
node of R,,; with more than one isolated setula (except in Neaerini and a few
Blondeliini). Parafrontals with the usual two pairs of proclinate orbital setae
in 9, without or with two pairs in g. Antenna not noticeably mucronate, outer
tip of last segment usually rounded. Wing venation of different form (slightly
similar in some Blondeliini). Second costal sector haired or bare below. Varied
complements of dc setae. [Not parasites of ee Acridoidea, a
possibly Phorocerosoma|

Subapical scutellar setae stronely convergent or Grossing a them: pe
(Text-figs 67 & 69). Second costal sector haired below (third sector also usually
haired ventrally on most of its length). Costal margin conspicuously broken
and incised at the apex of the subcosta (Sc) (Text-figs 86 & ty). Abdominal
Tr + 2 not excavate to its hind margin

Subapical scutellar setae parallel to each other or diverging fe: g. as in Text- ies a1).
Second costal sector bare below (except in Livophaga). Costal margin
inconspicuously broken and scarcely at all incised at the apex of Sc. Abdominal
Tr + 2 usually excavate to its hind margin (not in some Blondeliini or in
Hillomyia) , : é ; . : : .

Abdomen without discal setae on nany eeeite. Propleuron bare. Head not sexually
dimorphic, g with two pairs of proclinate orbital setae and with outer vertical
setae like the 9. Vein R,,, extensively setulose and F, usually also partly or
wholly setulose. Eyes bare. Hind tibia with three dorsal preapical setae
(pd preapical present in addition to the usual ad and d preapicals) (Text-fig 18)

Io

SIPHONINI (p. 80)

Abdomen with strong erect discal setae on tergites 3 to 5. Propleuron haired.
Head sexually dimorphic, frons much narrower in ¢ than Q and male head
without proclinate orbital setae or definite outer vertical setae. Vein Ry, ; with
one setula only or a small tuft of setulae confined to the basal node, R, bare.
Eyes haired (but hairing very short and inconspicuous in some forms). Hind
tibia with two dorsal preapical setae (the normal ad and d preapicals) (Text-fig. 19)

NEAERINI (p. 78)

Bend of vein M in the form of an open evenly rounded curve (as in Text-fig. 85) or
if shightly abrupt then forming a widely obtuse angle, always without trace of an
M, appendix or fold. Propleuron bare or haired. Infrasquamal hairs present
or absent. Abdominal Tr + 2 excavate or non-excavate to its hind margin.
Mid tibia with or without a submedian v seta ,

Bend of vein M abrupt and not forming a widely obtuse angle (JV changing direction
usually at about 90° at the actual bend), often provided with an M, appendix or

BLONDELIINI (p. 81)

78 R. W. CROSSKEY

fold in the wing surface (Text-fig. 87). Propleuron bare (except in Hillomyza).

Prosternum always haired or setulose. Infrasquamal hairs always absent.

Abdominal Tr + 2 excavate to its hind margin (except in Hillomyia). Mid tibia

always with a submedian v seta . : E : : : II
11 Lower calypter normal, outer edge not Beas loge eee Bend of vein M nearly

always with at least a trace of an M, appendix or at least a darkened fold in the

wing membrane continuing towards he wing margin (Text-fig. 87). Eyes bare

or haired. Three or four post dc setae. Barette vind haired only at its anterior

ends = i : EXORISTINI (p. 85)
— Lower calypter bent abraptly, downwards on its outer margin (except in

Mycteromyiella). Bend of vein M without trace of MM, appendix or fold. Eyes

densely haired. Four postdc setae. Barette fully fal . ETHILLINI (p. 87)

Tribe ACEMYINI

This is a small but nearly cosmopolitan tribe in which all the known hosts are
Orthoptera of the families Acrididae s.l. and Eumastacidae. The scutellar bristling
is distinctive among the Goniinae (Text-fig. 68) and comprises three pairs of very
strong setae, an apical crossed pair, a subapical pair set unusually far forwards
on the sides of the scutellum and a basal pair. Most forms have a long series of
proclinate orbital setae in both sexes and many have the outer tip of the third
antennal segment formed into a rather sharp point, these features helping towards
instant recognition of the group; the basal node of vein R,,; has only one extremely
strong setula on both surfaces of the wing, a feature that is relatively uncommon
in goniines, although found in the Neaerini. Only the single genus Ceracia Rondani
(syn. Myothyria Wulp) is known in Australia, and this is distinguished from other
acemyine genera by the following combination of characters: prosternum setulose;
head with proclinate orbital setae in both sexes; hind tibia without pv apical seta
(cf. Acemya Robineau-Desvoidy); humeral callus with three setae; cell R, closed
at or just before the wing margin (Text-fig. 88); two sternopleural setae; bend of
vein M rather abrupt and very remote from wing margin (Text-fig. 88); 2 + 3 dc
setae; vibrissae above the level of the epistomal margin; basicosta reddish yellow;
third antennal segment distinctly mucronate; head with inner eye margins strongly
diverging ventrally, parafacials not or hardly contracted at their lower ends.

Tribe NEAERINI

The Australian fauna contains several little known forms that are clearly very
closely allied to the genus Tongamyia Mesnil from Tonga and Fiji, and the
resemblance is indeed so complete that Voriella Malloch and Tongamyia ought
probably to be placed as synonyms (this is not considered further in the present
work as Voriella is the older name and would not have to be changed in the event
ofsynonymy). It is pertinent, however, that Mesnil (1960 : 649, footnote) associates
Tongamyia with the neaerines and the genus Neaera Robineau-Desvoidy, and
placement of Tongamyza in the Neaerini appears very reasonable on present evidence.
As Voriella is virtually indistinguishable from Tongamyia it, too, is here assigned

A

|
2

:

TACHINIDAE OF AUSTRALIA 79

to Neaerini. (The only noteworthy difference between Voriella and Tongamyia
lies in the vestiture of the hind coxa, as indicated in the key that follows.)

Voriella and its allies have the following main features in common. 4 without proclinate
orbital setae and with such fine straight reclinate orbital setae that they appear to form a
continuous row with the frontal setae; 2 with one pair of reclinate orbital setae (sometimes
slightly twisted outwards); vibrissae level with epistomal margin; frontal setae descending
far down on the parafacials (sometimes to a level below the mid-point) and in ¢ the rows
irregularly duplicated at the lower ends; upper occiput with some black setulae behind postocular
row; normally 2 + 3 dc setae (3 + 3 occurring, perhaps aberrantly); 1 + 3 ia setae; 2 stpl
setae; scutellum with only two pairs of strong setae, the basals and subapicals, the latter
meeting or crossing at their tips; propleuron haired; infrasquamal hairs absent; mid tibia with
a submedian v seta; mid tibia with one or two ad setae; hind tibia without $d preapical seta
and with or without a very small pu apical seta; second costal sector haired ventrally; costal
spine long; distance on vein M from bend to m-cu as great, or nearly so, as that from m-cu to
y-m (Text-fig. 86); costa of 3 often with long fine ventromarginal hair on basal part; basal node
of R,,, most often with only one strong setula above and below, sometimes with a hair tuft;
abdomen with Tr + 2 not excavate to hind margin and with very strong erect discal setae on
T3-T5.

In addition to the type-species, Malloch described two other Australian species
(viz. inconspicua and armiceps) that he placed in Voriella; neither correctly belongs
in the genus or in the Neaerini.

An undescribed species with haired parafacials has been seen from South Australia,
and another with the characters of Tongamyia but with 3 + 3 dc setae has been
seen from New South Wales. Both differ slightly from either Voriella or Tongamyia
and their generic placement is uncertain at present. They have been placed,
however, in the following key so that the differential features can be seen easily.
There are evidently more forms yet to be discovered in Australia in this interesting
and little-known complex. Probably when studied sufficiently it will be concluded
that all should be treated as congeneric, and the definition of Voriella widened
accordingly.

The BMNH collection contains an undescribed species of Neaerini from Queensland
that differs from all the other Australian Neaerini so far known by having the wing
cell R, exceedingly long-petiolate. This species is very small (length only about
2:5 mm) but differs from Tongamyia only by the petiolate cell R;. The species
will either require a new genus (if the long petiole is considered a sufficient generic
character) or will have to be placed in Tongamyia with the necessary widening of
the definition of the latter.

The affinities of the Neaerini appear to lie with the Siphonini, a tribe which
Herting (personal communication) considers should be assigned to the Tachininae.
Future work may well demonstrate that the neaerines also should be placed in
the Tachininae, and possibly also that the tribe should be merged with the
Eloceriini (=Helocerini of authors). In the latter event the correct tribal name
would be Eloceriini.

Key To AUSTRALIAN GENERA OF NEAERINI

I Wing with cell R; closed far before the wing margin, the petiole very long (longer than
either the second costal sector or M,) [Queensland] . . | Undescribed sp., ? gen. n.

80 R. W. CROSSKEY

— Wing with cell R,; open or at most only just closed at the wing margin. - : 2
2 Parafacials with a row of strong black hairs or setulae (curving downwards). Basal
node of vein R, , , with several long hairs or fine setulae on upper and lower surfaces,
those of lower surface often forming a definite tuft [South Australia]
Undescribed sp., ? gen. n.
— Parafacials bare (apart from the strongly descending frontal setae). Basal node of
vein R,,; usually with only one long strong setula on upper and lower surfaces,

at most only two minute hairs in addition to the main setula : 3
3. Mid tibia with one submedian ad seta. Hind coxa bare on posterodorsal eaarpint

2 + 3 dc setae [Fiji, Tonga, Samoa] ‘ ; TONGAMYIA Mesnil
— Mid tibia with two ad setae (a definite small ad oe nea just basad of the main

one). Hind coxa bare or setulose on posterodorsal margin. 2 + 3 or 3 + 3 dc

setae [Australia] . : : E : : c 4

4 Hind coxa with black setulae on the peste cael fae 2 + 3 de setae. Fore
tibia with two pv setae (basal one weak) [widespread eastern Australia]
VORIELLA Malloch
— Hind coxa bare on the posterodorsal margin. 3 + 3 dc setae (a third prst dc seta
present between the main two and almost equally strong: ? aberrant). Fore tibia
with one pu seta [may not be constant when more material known] [New South
Wales] : : - : ; . : : : : Undescribed sp.

Tribe SIPHONINI
(Actiini)

The cosmopolitan tribe Siphonini is the most distinctive in the Goniinae, and the Australian
forms can be instantly recognized by their characteristic facies. This derives primarily from
the following features: size very small (length about 3-4 mm, smallest Australian Tachinidae
on average); head not sexually dimorphic (the sexes therefore rather difficult to recognize as
both males and females have an equally broad frons, two pairs of proclinate orbital setae and
strong outer vertical setae); scutellum with subapical setae convergent and usually crossing
towards the tips (Text-fig. 67); pre-alar seta minute; legs rather short and with rather stiff
setae (hind tibia unusual in having three strong preapical setae on upper surface —1.e. ad, d
and pd preapicals — as shown in Text-fig. 18); abdomen without discal setae on any segment
and with Tr + 2 excavate only on its anterior half; wings short and broad with costal margin
deeply incised at apex of Sc and with bend of vein M unusually remote from cross-vein m-—cu
(Text-fig. 89); wing veins more extensively setulose than in other goniines.

The Australian distribution includes Tasmania, and there are three genera
represented in the area. The most notable absentee is the genus Szphona Meigen,
which although represented by at least one species in New Guinea has not yet been
found in Australia. This genus is not included in the key as, if found later in
Australia, it will be immediately recognisable by the extremely long slender angled
proboscis that is at least as long as the head height (all genera known in Australia
have a short inconspicuous proboscis).

It is necessary to note here some changes in nomenclature of Australian siphonines.
Crosskey (1966b) applied the generic name Strobliomyia Townsend to several
Australian species, using the nomenclature given by Mesnil (1963). Mesnil noted
two older names for Strobliomyia, viz. Herbstua Robineau-Desvoidy and Peribaea
Robineau-Desvoidy, but considered that they were both preoccupied and therefore
that Strobliomyia was the valid name. Mesnil was right about Herbstia Robineau-

TACHINIDAE OF AUSTRALIA 81

Desvoidy which is preoccupied by Herbstia Edwards but wrong about Peribaea
which is not preoccupied. The valid name for the genus previously called
Strobliomyia is now, therefore, Peribaea. Mesnil (1963 : 803) placed the names
Schizoceromyia Townsend and Schizactiana Curran as synonyms of Strobliomyia
(= Peribaea) but these synonymies are in error; the type-species of Schizoceromyia
and Schizactiana have been examined, and neither has the strong downwardly
directed prostigmatic seta that characterises Peribaea; on the contrary, both
type-species have all the essential characters of Ceromya Robineau-Desvoidy, and
the names Schizoceromyia and Schizactiana are therefore moved into new synonymy
with Ceromya.

Some species of Peribaea have the third antennal segment deeply cleft in the
male, and a bifid male third antennal segment occurs in fergusoni Bezzi and valida
Curran (the type-species of Schizoceromyia and Schizactiana respectively). Mesnil’s
placement of Schizoceromyia and Schizactiana in synonymy with Peribaea (as
Strobliomyia) was evidently due to this feature —as the type-species were known
to him only from descriptions, which did not mention the prostigmatic setae.
It is now clear that forms with bifid or ramose male third antennal segments occur
in both Peribaea and Ceromya —and indeed also in Actia (in which A. baldwini
is one such species). Cleaving of the third antennal segment is extremely rare in
the Australian tachinid fauna, and these siphonines are the only described Australian
forms in which such a characteristic occurs.

The Siphonini are here placed in their traditional position, but there are reasons
for thinking that they would be better placed in the Tachininae—notably certain
features in the larval morphology (Herting, personal communication).

KeEy TO AUSTRALIAN GENERA OF SIPHONINI

I Thorax with two strong subequal prostigmatic setae, one directed upwards and the
other downwards (Text-fig. 8). Wing with sixth vein extending to wing margin.
Mid tibia with one strong ad seta . : : . PERIBAEA Robineau-Desvoidy
-— Thorax with one strong prostigmatic seta, directed upwards. Wing with sixth vein
usually stopping short of wing margin (occasionally traceable to peg as weak
fold). Mid tibia with or without anadseta . : 2
2 Sternopleuron with a regular row of fine hairs in front of the mid coxa (Text- fie. I 4)
Lowermost sternopleural seta very weak and placed about as far from the mid
coxal base as from the upper anterior thee seta [condition not known for
: all included species] : - . ACTIA Robineau-Desvoidy
Sternopleuron completely bare. laterally in front of the mid coxa. Lowermost
Pas ccn) seta either very strong and placed very near the mid coxal base or
(in feygusont) undeveloped so that there are only two sternopleural setae [condition
not known for all included species} : : . CEROMYA Robineau-Desvoidy
| -
|
|

Tribe BLONDELIINI

This tribe contains a very large number of genera each with relatively few species.
| There is great diversity of body form and chaetotaxy within the group, but it is
| usually possible to recognize the blondeliines reliably among the goniines by the

possession of three main features in combination, viz. pre-alar seta small or minute
F

82 R. W. CROSSKEY

(weaker than first post ia seta), subapical scutellar setae extremely strong and
divergent (with the apicals very weak and often absent) (Text-fig. 71), and the bend
of vein M forming an evenly rounded curve (Text-fig. 85). Some forms occur that
do not completely fit with these criteria, but they are nevertheless very helpful in
practical recognition of the group; some forms may have fairly well developed
apical scutellar setae, and a very few (such as the Australian Paropsivora) have a
rather abruptly angulate vein M. The tribe is nearly cosmopolitan, and is well
represented in Australia and Tasmania (whence many undescribed forms exist in
collections in addition to the identifiable genera and species). The hosts of many
blondeliines are beetles or sawfly larvae, insect groups that much less commonly
provide hosts for the other tribes of Goniinae.

Some of the Blondeliini with black coloration and slender elongate bodies and
legs are strongly reminiscent of the Minthoini and it seems possible that there
is a much closer affinity between blondeliines and minthoines than has been supposed
or that some forms are erroneously classified in the Blondeliini (e.g. the tropical
genus Eophyllophila Townsend). This genus occurs in New Guinea but has not
been found in Australia, but black-and-yellow forms of the genus Trigonospila
(which is evidently a close relative of Eophyllophila) are found in Australia. Other
blondeliines, such as Froggattumyia, have very much the facies of Sturmiini and
the existence of such forms as well as minthoine-like forms in the same tribe makes
it difficult for the non-specialist to recognize the tribe on general appearance; but
at least virtually all the members, whatever their naked-eye facies, share the three
main characteristics already noted.

Another noteworthy character found in many blondeliines is hairing on the
propleuron. This occurs in several quite different groups of blondeliines, but
almost never in other tribes of Goniinae (in which the propleuron is bare except in
Hillomyia). It is therefore a useful rule-of-thumb when identifying Australian
tachinids that any gonine specimen with a haired propleuron belongs in the
Blondeliini. The Australian fauna is rich in forms with haired propleuron and
contains one notoriously difficult complex in which the supposed genera merge rather
imperceptibly into one another. This complex includes Anagonia and related forms
and is currently under study by Dr Donald Colless.

Very nearly all species of the enormous subfamily Goniinae have the prosternum
haired or setulose (or at least one pair of prosternal setulae) but some blondeliines
are atypical members of the Goniinae and have the prosternum totally bare. In
Australia such forms include the genera Tvigonospila and Zosteromeigenia, which
(excepting rare aberrant specimens of other genera) are the only Australian Goniinae,
apart from the sturmiine genus Blepharella, to have a bare prosternum.

About half of the genera of Blondeliini so far known from Australia are endemic,
but this proportion is likely to rise when the fauna is better known. The non-endemic
genera occur widely in Eurasia and at least two (Compsilura and Trigonospila)
are found also in tropical Africa. Lixophaga is mainly a New World genus but has
at least two species (one undescribed at present) in New Guinea; the described
species, L. sphenophori, was introduced into Queensland and is apparently established
there. The New Guinea blondeliines have not been worked out, but the genera

TACHINIDAE OF AUSTRALIA 83

Eophyllophila Townsend and Prodegeeria Brauer & Bergenstamm certainly occur
in the territory and may perhaps be found ultimately in Queensland.

The genus Bactromyiella is very anomalous, but may perhaps be a blondeliine.
It has a small pra seta (smaller than the first post 1a seta) and the bend of vein M
is evenly rounded as in typical Blondeliini, but the subapical scutellar setae are
parallel (not strongly divergent as is usual in blondeliines); the facies of Bactromyvella
is not especially blondeliine, but is more that of a winthemiine or eryciine. The
genus is kept tentatively in the Eryciini but is included also in the following key
to blondeliine genera.

Kery TO AUSTRALIAN GENERA OF BLONDELIINI

t Mid tibia without a submedian v seta 5 - , 4 : ; 4 : 2
Mid tibia with a submedian v seta. 84

2 3+ 4 dc setae. Prosternum haired. Scutellum with subparallel subapical setae

and with apical setae. Intermediate abdominal tergites without discal setae.

6 with velvety black thorax contrasting with bright orange or orange-red abdomen
(except Tr + 2 black) . ; [BACTROMYIELLA Mesnil, tribe Eryciini]

— 2+ 3 de setae. Prosternum bare. Scutellum with widely diverging subapical

setae and without apical setae. Intermediate abdominal tergites with strong
erect discal setae. d without such coloration : TRIGONOSPILA Pokorny

3 2-+ 2 dc setae (sometimes a very small additional post dc setula between the two

main setae). Second supra-alar seta absent. Humeral callus with two setae.

Wing with distance on vein M from the bend to m—cu as great as that between

m-—cu and v-m. Basal node of R,,, bare or with one minute hair. Two
sternopleural setae. Abdomen with Tr + 2 not excavate to its hind margin. 4

More than 2 + 2 dc setae (2 + 3 or 3 + 3 or 3 + 4, occasionally 2 + 4). Second

supra-alar seta present. Humeral callus with three or more setae. Wing with

distance on vein M from the bend to m-—cu conspicuously less than that from

m—cu to y-m. Basal node of R,,, with at least two hairs or setulae. Two or

three sternopleural setae. Abdomen with T1 + 2 excavate to its hind margin
(except in Livophaga and Lecanipa) : 5

4 Propleuron bare. 4 with one pair of strong areas discal setae on each sedoniinal

tergite from T3 to T5, 2 without abdominal discal setae. 4 without proclinate

orbital setae. with ovipositor in form of a long strong downcurved hook easily

seen in profile. Q hind coxa with en setae modified into short blunt
black pegs : : . MEDINODEXIA Townsend

- Propleuron sparsely haired: Both. sexes Grithout abdominal discal setae. G with

two pairs of proclinate orbital setae (like the 9). Q with very short broad flattened

ovipositor that is not visible in profile. 9 hind coxa with apicoventral setae not
so modified : : : MONOLEPTOPHAGA Baranov

5 Propleuron bare. Vibrissae inserted anoae on a level with the epistomal margin.

Parafacials bare (except in Pareupogona). ae sari hairs sometimes
present ‘ 6

— Propleuron haired. Vibrissae. usually inserted at a eae cisuinetly above the

epistomal margin. Parafacials usually entirely haired or haired at upper ends,
seldom entirely bare. Infrasquamal hairs absent . : 10

6 Fore tibia with two pu setae. Intermediate abdominal tergites with discal setae.

Second costal sector bare ventrally. Hind tibia without a pd preapical seta.

Humeral callus with the three main setae standing in a straight line es in
Lecanipa). 4 without proclinate orbital setae : Ff

— Fore tibia with one pv seta. Intermediate abdominal tergites without ‘discal

84

It

I2

13

14

Re W. CROSSKEY

setae. Second costal sector haired ventrally. Hind tibia with a distinct pd

preapical seta in addition to the normal ad and d preapicals. Humeral callus

with three setae standing in a triangle. ¢@ with two pairs of proclinate orbital

setae (like the 9) é LIXOPHAGA Townsend
Eyes bare. Parafacials aoc and Spanely heed Facial ridges finely setose on

lower two-fifths or half. Mid tibia with two ad setae PAREUPOGONA Townsend
Eyes densely haired. Parafacials bare. Facial ridges with strong downcurved

setae on most of their height. Mid tibia with one or two ad setae . 8
Mid tibia with one submedian ad seta. 3+ 4 dc setae. Abdominal Tx ig 2

excavate to its hind margin. Humeral callus with the three main setae standing

in line. Infrasquamal hairs absent - : . 3 ; : : 9
Mid tibia with two ad setae. 3 + 3 dc oe (supernumerary fourth post dc

occasionally present between the normal first and second post dc setae).

Abdominal Ti + 2 with excavation clearly not reaching hind margin. Humeral

callus with the three main setae standing in a triangle. Infrasquamal hairs

present : : . LECANIPA Rondani
Three sternopleural setae. Tergites of fe) padoned tone ed to form a mid ventral

abdominal keel bearing stubby spinules. 9 with ovipositor in form of a strong

downcurved hook-like piercer : - : COMPSILURA Bouché
Two sternopleural setae. Tergites of 9 abdomen normal. 9 without externally

visible piercing ovipositor . : : : Undetermined genus [? gen. n.]
2+ 3 dc setae. Two sternopleural setae. Eyes bare. Parafacials fully haired.

Intermediate abdominal tergites with discal setae . . DELTOMYZA Malloch

Dorsocentral setae 3 + 4 or 3+ 3 (rarely 2+ 4). Three sternopleural setae
(except sometimes in Zosteromeigenia). Eyes bare or haired. Parafacials haired
or bare, or partially haired. Intermediate abdominal tergites without discal
setae (except in Paropsivora) : II
Three post dc setae. Abdomen of 3 withie a fete of long dense hair ¢ on éach side
of T5 (last visible tergite). Parafacials mainly or totally bare, if haired then

hairing confined to area immediately below lowest frontal setae : : : 12
Four post de setae. Abdomen of ¢ without dense hair fascicles on T5. Parafacials
completely haired or at least haired on uppermost quarter or third . : 13

Fore tibia with two pu setae. Mid tibia withtwoadsetae. Intermediate abdomaal
tergites with discal setae. Humeral callus with four setae (basal row of three
in which median one weakest, and a fourth seta set forwards of the basal row
and stvongey than middle seta of the basal row). Bend of vein M strongly
abrupt. Eyes haired. Prosternum haired . : . PAROPSIVORA Malloch
Fore tibia with one pu seta. Mid tibia with one ad seta. Intermediate abdominal
tergites without discal setae. Humeral callus with only two strong setae,
sometimes a rather weak third seta. Bend of vein M not strongly abrupt.
Eyes bare [haired in an undescribed New Guinea species]. Prosternum bare
[haired in undescribed New Guinea species] . ZOSTEROMEIGENIA Townsend
Facial ridges bare (except for the usual few setulae immediately above the vibrissae).
Eyes bare or haired. 9 with the usual well developed outer vertical setae.
Vibrissae conspicuously above the level of the re) margin (except in
Zenargomyia) : ; E : : 14
Facial ridges strongly Seonlese up most ‘of their height. Eyes densely haired.
2 without outer vertical setae [character in need of confirmation as very few
Pilimyia females seen]. Vibrissae inserted only slightly above the level of the
epistomal margin : . PILIMYIA Malloch
Vibrissae inserted only very slightly eae the level of the epistomal margin.
Eyes bare. Parafacials bare except on the uppermost quarter or third (Text-fig.
49). Ocellar setae very strong. Apical scutellar setae rather strong and crossed
ZENARGOMYIA Crosskey

TACHINIDAE OF AUSTRALIA 85

— Vibrissae inserted well above the level of the epistomal margin. Eyes bare or
haired. Parafacials completely haired or haired at least on the upper halves.
Ocellar setae usually absent or weak, sometimes moderately strong in9. Apical

scutellar setae usually very weak and hair-like, sometimes absent, seldom well
developed . P : 5

15 Facial regions of head with yellow or orange ground colour and bright yellow
pollinosity. Eyes bare. Basicosta reddish yellow. Legs usually all or mainly
reddish yellow, at least tarsi reddish yellow. Haired areas of parafacials not
nearly contiguous with haired areas of genal dilations (therefore a conspicuous
bare area between haired part of parafacial and the gena). On average larger
forms, length 7-12 mm, often with yellow to golden orange hair on sides of thorax
and sometimes also the abdomen . ; : . FROGGATTIMYIA Townsend

— Facial regions of head with rather indeterminate ground colour, not bright yellow
and if pale usually rather reddish, pollinosity greyish white to very pale yellowish.
Eyes bare or haired, often densely haired. Basicosta nearly always dark brownish
or blackish brown. Legs brownish black, sometimes with reddish or reddish
yellow tibiae, tarsi dark (except for reddish yellow fore tarsi in male of A.
scutellata). Haired areas of parafacials nearly contiguous with haired areas of
genal dilations (only a very narrow bare gap between parafacial and genal
hairing). On average smaller forms, length 5-9 mm, without pale hair on thorax
or abdomen . : . : ; . MANAGONIA Brauer & Bergenstamm

[Note: no really tangible distinction has been found between Froggattimyia
and Anagonia|

Tribe EXORISTINI

The exoristines are moderately well represented in Australia and Tasmania,
seven genera being known from the area. None is endemic, and several of them
occur widely in the Oriental Region (Eozenillia, Austrophorocera) or in most of
the zoogeographical regions (Exorista, Stomatomyia); Hillomyia (= Hillia Malloch,
preocc.) is known from only a few specimens but is perhaps an essentially Papuan
genus that has reached the Northern Territory of Australia.

The tribe is reasonably distinctive, possessing the combination of small pre-alar
seta plus non-convergent subapical scutellar setae plus abruptly angulate bend to
vein M (the bend normally accompanied by at least a trace of a fold in the position
of M, continuing towards the wing margin, Text-fig. 87); the genera found in
Australia have unusually dependable characters, and most are considerably easier

: to identify reliably than the genera in most other goniine tribes. Some features
common to all the Australian exoristines may be usefully noted as follows:
prosternum always haired; male always without proclinate orbital setae; mid
tibia with a submedian v seta; mid tibia with two or more ad setae; hind tibia
always without #d preapical seta; second costal sector bare ventrally; infrasquamal

| hairs absent; parafacials never completely haired.

|
|
| Key To AUSTRALIAN GENERA OF EXORISTINI

| I 2+ 3 dc setae. Wing with cell R, closed before the wing margin and with a

1 distinct petiole. Distance on vein M from bend to m-—cu as great as or greater

than that from m-—cu to v—-m. Scutellum without definite preapical setae.
Humeral callus with two setae. Eyes bare . ; . : c : : 2

86

R. W. CROSSKEY

3 +3 0r 3+ 4 dcsetae. Wing with cell R,; not closed before the wing margin, at
least narrowly open. Distance on vein M from bend to m—cu much less than
that from m—cu to v-m (except in one species of Stomatomyia). Scutellum with
a pair of preapical setae. Humeral callus with three or four setae. Eyes bare
or haired . g 5 ; : : : : 3
Abdomen with Tr + 2 eee to its hind meee Vein R, setulose. Fore tibia
with two pu setae. Propleuron bare. Scutellum with strong upwardly directed
apical setae. Upper occiput without black setulae behind the postocular row.
Abdomen with conspicuous white-pollinose bands on T3 to T5 . CHAETORIA Becker
Abdomen with excavation of Tr + 2 not reaching the hind margin of the tergite.
Vein R, bare. Fore tibia with one pu seta. Propleuron with a few long white
hairs [careful examination needed]. Apical scutellar setae almost absent,
represented by a pair of fine hairs. Upper occiput with a few fine black setulae
behind the postocular row. Abdomen uniformly black and rather shining
HILLOMYIA Crosskey
3 + 3 dcsetae. Last section of vein Cu, (from m—cu to wing margin) very long,
more than half as long as the penultimate section and very much longer than
m-—cu; m-—cu joining Cu, at an oblique angle. Apical scutellar setae directed
very strongly upwards. Costal spine well developed, usually very easily dis-
tinguishable from marginal costal setulae and as long as cross-vein y—m or longer . 4
3 + 4 dc setae (except in one unidentified sp. from Northern Territory). Last
section of vein Cu, normal, shorter than or subequal in length to m—cu and
much less than half as long as the penultimate section; m—cu joining Cu, approxi-
mately at a right-angle. Apical scutellar setae nearly horizontal or directed
only slightly upwards (except in some Evxorista). Costal spine not developed,
indistinguishable from marginal costal setulae 2 ‘ :
Abdomen without discal setae on intermediate tergites (T3 soe T4). Palpi well
developed. @ with upper pair of reclinate orbital setae showing at most only a
slight outward twist towards the eyes. Apical scutellar setae weak, very much
smaller than the lateral scutellar setae. 4 with or without outer vertical setae,
if present then obviously weaker than ocellar setae
STOMATOMYIA Brauer & Bergenstamm
Abdomen with two or more very strong erect discal setae on each intermediate
tergite, accompanied by numerous smaller stiff erect setulae. Palpi absent or
almost so (at most represented by a minute papilla bearing a long apical hair)
(well developed in an undescribed species). @ with the upper pair of reclinate
orbital setae enormously strong and directed outwards over the eyes (simulating
prevertical setae). Apical scutellar setae extremely strong, almost as large as
lateral scutellar setae. G with very strong outer vertical setae subequal in size
to ocellar setae . : : : SPOGGOSIA Rondani*
Facial ridges armed with strong downeurved setae on most of their height. Eyes
densely haired. Robust forms with short inconspicuous M, fold extending
from bend of vein M; hypopygium of ¢ never with golden hair : 6
Facial ridges mainly bare, only with weak hairs and setulae immediately abeee the
vibrissae which extend at most only for one-third of the height of the facial
ridges. Eyes bare or haired. More slender forms with rather narrow tapering
abdomen and with a long conspicuous M, fold in the wing extending from the
bend of M; ¢ often with dense golden hair on the hypopygium (visible in situ)
EXORISTA Meigen
3 + 4 dc setae. Ocellar setae absent or very weak and wiry. Frons and thoracic
dorsum not golden : ; F z : : : : : . : 7
3 + 3 dc setae (? aberrant). Ocellar setae very strong (much larger than upper pair
of reclinate orbital setae). Frons and thoracic dorsum conspicuously golden to
naked eye . : : : : ; : : Undetermined genus & species

On

* See Appendix, p. 209.

TACHINIDAE OF AUSTRALIA 87

[Two specimens (¢ and 9) are in BMNH, London, collection that run out here.
They are from Northern Territory. Superficially they much resemble the genus
Phorinia Robineau-Desvoidy and should perhaps be assigned to it. The presence
of only three post dc setae may be an aberration, as four would be expected from
the total facies. ]
7 Interfrontal area at mid point subequal in width to parafrontal or at least two-thirds
as wide. Second aristal segment not more than twice as long as wide
AUSTROPHOROCERA Townsend
— Interfrontal area at mid point not more than half as wide as parafrontal in most
specimens, occasionally up to two-thirds as wide. Second aristal segment
about 2-5—4 times as long as wide ; : . EOZENILLIA Townsend
[This is the least satisfactorily distinguishable genus in the Australian exoristine
fauna. The one included species, E. vemota, is undoubtedly congeneric with
Eozenillia type-species and is therefore assigned to Eozenillia. Distinctions
between Eozenillia and Austrophorocera are very intangible, and probably the
former should be synonymised with the latter. ]

Tribe ETHILLINI

This small group was originally defined by Mesnil (1944) to include an aggregate
of forms possessing a facies much like that of the Winthemiini or Sturmiini but
differing from these in having a small pre-alar seta and by having the outer edge
of the lower calypter conspicuously bent downwards. These characteristics
undoubtedly help to define what appears to be a natural group, although some
species of undoubted sturmiines and carceliines have a similar ‘downbent’ calypter.
Conversely some forms such as Mycteromyiella in which the lower calypter is not
markedly bent down on its outer margin seem to belong in the Ethillini. The
undescribed New Guinea fauna contains several interesting forms which to a large
extent bridge the gap between Phorocerosoma (placed by Mesnil in the ethillines)
and Mycteromyiella (placed by Mesnil in the sturmiines) and it seems most
appropriate —on present knowledge at least—to assign Mycteromyiella to the
Ethillini.

The known hosts of Phorocerosoma are acridid grasshoppers and the known
hosts of Mycteromyiella are species of Mantodea, but host records have not yet
been obtained in Australia itself. The tribal distribution includes Tasmania
as well as continental Australia. It is important to note that the characters given
in the following key for Phorocerosoma apply to the single species known to occur
in Australia, but do not necessarily fit extra-limital species of the genus. The
generic identities of most of the New Guinea ethillines are too uncertain at present
for these forms to be covered in the key.

Kry To AUSTRALIAN GENERA OF ETHILLINI

I Inner vertical setae converging and crossing before their apices. Abdominal
Ti + 2 and T3 each with two pairs of very long strong erect median marginal
setae. Abdomen without discal setae on intermediate tergites. Humeral
callus with the three main setae standing in a triangle. 9 with a pair of strong
prevertical setae that curve outwards over the eyes and without outer vertical
setae. Pre-alar seta longer than first postiaseta . PHOROCEROSOMA Townsend

88 RR. W. CROSSE

— Inner vertical setae subparallel. Abdominal Tr + 2 and T3 each with one pair of
median marginal setae (those of Ti + 2 sometimes hair-like or virtually absent).
Abdomen usually with one or two distinct discal setae on one or both intermediate
tergites. Humeral callus with three main setae standing either in a straight
line or in a triangle. Q without outwardly directed prevertical setae and with
the usual pair of outer vertical setae. Pre-alar seta very small, shorter than
first post ia seta . 3 : : ¢

2 Humeral callus with the three main setae Sear ina straight ae. Lower calypter
with the outer edge distinctly bent downwards. Arista with second segment
elongate, about 4—5 times as long as broad. Head with two pairs of strong
reclinate orbital setae. Mid tibia with one ad seta. Vibrissae level with
epistomal margin : : ‘ ETHILLA Robineau-Desvoidy

— Humeral callus with the ee main ‘sete eden in a distinct triangle. Lower
calypter not noticeably bent downwards. Arista with second segment not
elongate, only about as long as broad. Head without definite reclinate orbital
setae (at least in g, probably one pair in 9). Mid tibia with two ad setae.
Vibrissae inserted well above level of epistomal margin

MYCTEROMYIELLA Mesnil

nN

Tribe WINTHEMIINI

This group is very closely allied to the Carceliini and the Sturmiuini and its
recognition as a tribe is only doubtfully justified. Townsend (1941) placed Winthemia
and its allies in the Sturmiini, but recent workers universally treat the winthemiines
as a valid tribe and this course is followed for the present work. The arrangement
of the humeral setae in Winthemia is rather distinctive, there being (in males at
least) five setae in which three are strong and are spaced in a subtriangular arrange-
ment; in this respect Winthemia differs from the Sturmiini in the strict sense in
which the humerals are four arranged as three in a straight line and one set forwards.
Mesnil (1944) used the presence of five humeral setae as a key character for recognition
of the winthemiines, but in reality the character is poor — often in other winthemiine
genera than Winthemia itself (e.g. in Nemorilla) five setae may not be definitely
developed in males and are typically not developed in females (which usually have
four humerals).

The winthemiines are best recognized among the Australian Goniinae by the following
features in combination (though some Sturmiini and some Carceliini may show some of the
features together): eyes densely haired; males without definite reclinate orbital setae; hind
tibia (especially in males) with a well developed close-set ad fringe; barette completely haired
(or nearly so); scutellum with strong crossed horizontal apical setae and with subapical setae
inserted widely apart (their bases as distant from each other as base of either from the basal
seta of the same side, or even more distantly separated than this) (as in Carcelia, Text-fig. 72).
A noteworthy feature of several forms is the presence of pale yellow hair on the prosternum
instead of the usual black hairs or setulae.

There is close superficial resemblance between Winthemiini and Ethillini, and
as the genus M ycteromyiella (Ethillini) could easily be confused with the winthemiines
it is included in the following key.

Mesnil (19494 : 80) treated Crypsina as a subgenus of Winthemia but it is here
preferred to revert to its original generic status. This course maintains Winthemia
as a more homogeneous genus, though — certainly — Crypsina is very closely allied.

TACHINIDAE OF AUSTRALIA 89

Kry To AUSTRALIAN GENERA OF WINTHEMIINI

1 Parafacials haired on their whole extent . : : - - é ‘ : 2
Parafacials bare i : : : : : : : - : :
2 Mid tibia with 2 or 3 strong ad setae. Vibrissae set high above epistomal margin
(distance between level of their insertions and the margin of the epistome at
least as great as width of antenna). Vibrissal insertions unusually close to each
other, distance between vibrissae less than or not noticeably greater than distance
between a vibrissa and the eye. Antennae bright orange
CRYPSINA Brauer & Bergenstamm
— Mid tibia with one submedian ad seta. Vibrissae not or only a little above level of
epistomal margin. Vibrissal insertions normal, distance between them much
greater than distance between vibrissal base and the eye. Antennae largely
blackish brown, or if extensively reddish orange then normally suffused with
brown at least along forward edges of third segment
WINTHEMIA Robineau-Desvoidy
3 Three sternopleural setae (2 + 1). Scutellum without lateral setae. Vibrissae
inserted far above epistomal margin (by a distance greater than width of third
antennal segment). Pre-alar seta weak, shorter than first post ia seta
([MYCTEROMYIELLA Mesnil, tribe Ethillini]
— Two sternopleural setae (1 + 1). Scutellum with lateral setae. Vibrissae level
with or only slightly above epistomal margin. Pre-alar seta strong, longer than
first post ia seta . ; 2 ; é - : : NEMORILLA Rondani

Tribe CARCELIINI

As it is understood at present this tribe contains those Goniinae that have a
large pre-alar seta and the genal region exceedingly narrow (the eyes being relatively
so large that they occupy most of the side of the head with consequent reduction
of the gena to a narrow strip below the eye that is narrower than the profrons)
(Text-fig. 48). The Anacamptomyiini are similar in head facies, but are treated
as distinct because of their unusual hosts (see under treatment of Anacamptomyiini
below). Some forms closely resemble the Winthemiini because of their densely
haired eyes and arrangement of scutellar setae; the genus Carcelimyia is especially
like the winthemiines because the barette is completely haired.

On present evidence the tribe appears to be rather poorly represented in
Australia, in contrast to its rich development in the Oriental Region. It is
probable, though, that other carceliine genera than those already known will
be found eventually in Queensland; this is especially likely because several genera
(e.g. Thecocarcelia) are known from New Guinea, and there seems to be no reason
why they should not occur also in northern Queensland. The distribution includes
Tasmania, from which one species is known (Carcelia tasmanica Robineau-Desvoidy).

KeEy To AUSTRALIAN GENERA OF CARCELIINI

I Four sternopleural setae. Eyes bare. Antennae exceptionally long, nearly reaching
epistome. Q with flat shining sclerotized ovipositor (visible externally)
THECOCA RCELIA Townsend
— Two sternopleural setae. Eyes bare or haired. Antennae not exceptionally heavy,
normally not reaching as far as the epistome. Y without such ovipositor : 2

go RW. CROSSKE ¥

2 Eyes bare. Apical scutellar setae very small, directed upwards and either not
meeting or crossing at tips. Subapical scutellar setae not conspicuously distant
from each other, their bases separated by a distance not greater than that
between a subapical seta and its corresponding basal seta. Upper occiput
with some fine black setulae behind postocular row
ARGYROPHYLAX Brauer & Bergenstamm
— Eyes haired. Apical scutellar setae strong, crossed and horizontal. Subapical
scutellar setae very widely separated, distance between bases much greater
than that between base of a subapical seta and its corresponding basal seta
(Text-fig. 72). Upper occiput without black setulae behind postocular row ; 3
3 Dorsocentral setae 3 + 4. Scutellum with extremely strong apical setae that are
at least as strong as, and usually larger than, the lateral setae. Hairing of
eyes long and dense, very conspicuous. Barette normally with hairing, at
least at anteriorend. Wingshyaline. Not conspicuously black forms :
— Dorsocentral setae 3 + 3 (four post dc setae in occasional specimens). Scutellum
with apical setae normally shorter and weaker than lateral setae. Hairing of
eyes rather short and usually rather sparse, not so immediately conspicuous.
Barette totally bare. Wings suffused with brown anterobasally. Conspicuously
black forms with shining abdomen which shows bright silver pollinose areas on
each side of T3-Ts5 (these especially noticeable on T4, at least in Q)
ARGYROTHELAIRA Townsend
4 Facial ridges bare. Barette when haired with hairing confined to anterior half
(except sometimes in specimens of Carcelia s. str.) . CARCELIA Robineau-Desvoidy
— Facial ridges closely setulose up most of their height (Text-fig. 48). Barette
completely haired : ‘ : : : : : CARCELIMYIA Mesnil

KEY TO AUSTRALIAN SUBGENERA OF CARCELIA

1 Mid tibia with a submedian v seta. Ocellar setae absent or very small, fine and
wiry. 4 with one pair of reclinate orbital setae, 9 usually with two pairs (specimens
with two such setae on one side and one on the other occur). Hind coxa bare
or setulose posterodorsally . c subgenus CARCELIA Robineau-Desvoidy
— Mid tibia without a v seta. Ocellar setae very strong (except in muvina where
absent or hair-like). Both sexes with two pairs of reclinate orbital setae. Hind
coxa entirely bare posterodorsally. : . subgenus SENOMETOPIA Macquart

Tribe ANACAMPTOMYIINI

Townsend (1940) recognized a tribe for Anacamptomyia and its allies and
placed it near Compsilurini (i.e. Blondeliini), but Mesnil (1944) included the
anacamptomyiines among the carceliines. On adult morphology Mesnil’s placement
seems to be more appropriate than Townsend’s, and in fact it is difficult to find
characters that satisfactorily differentiate the Anacamptomyiini from the Carceliini.
Nevertheless the Anacamptomyiini is here treated as a valid tribe because of its
biology: the members of the tribe are unique amongst the Goniinae in being
parasites of paper-making wasps (such as Belonogaster and Polistes), and this
host-relationship is so exceptional that it appears justified to give it considerable
weight in assessing whether or not to treat the flies involved as carceliines or not.
It appears possible that the resemblances between Carceliini and Anacamptomyiini
are mainly convergent.

TACHINIDAE OF AUSTRALIA QI

Mesnil (1944 : 27) synonymized Malloch’s Australian genus Vespivora with the
African genus Anacamptomyia Bischof. This synonymy is accepted here, but it
is pointed out that the only Australian species, viz. nigriventris (the type-species
of Vespivora) differs in several respects from typical Anacamptomyia; for example
the eyes are bare, there are no discal setae on the intermediate abdominal
segments, the median marginal setae of T,,. are weak or absent and there are
only two # setae on the mid tibia. On the other hand, the general facies of
Vespivora, the setulose facial ridges and the upwardly directed and crossed apical
scutellar setae, and the tendency towards reclinate uppermost frontal setae,
conform with Anacamptomyia (though the apical scutellars are much weaker than
the enormous apicals in the African Anacamptomyia species).

The genus Koralliomyia is tentatively placed in this tribe because of the
Hymenopterous hosts and the anacamptomyiine-facies (head profile as Text-fig. 52).

The species in this tribe are rather unusual amongst goniines for the pallid
reddish or reddish yellow colouring of most specimens. In all the Australian
species the basicosta is clear pale yellow-orange and the legs (including the tarsi)
are usually all reddish yellow (sometimes the femora partially dark brown in
nigriventris). So far the Australian distribution is only known to include Queensland
and New South Wales. One genus, Euvespivora Baranov, has very recently been
discovered in Hawaii (Oahu).

Biologically the Anacamptomyiini are an Old World counterpart to the small
tribe Ophirionini of the Neotropical region, members of which also parasitize social
Vespidae (especially the genus Polybia Lepeletier).

Kry To AUSTRALIAN GENERA OF ANACAMPTOMYIINI

I Propleural seta present. Parafrontals meeting in mid-line of frons, interfrontal
area therefore obliterated or almost so. Antennae very small, antennal axis below
mid-eye level. Mid tibia with at least two strong ad setae. [Parasite of Ropalidia
and Polistes] . ‘ : ; ; ; , KORALLIOMYIA Mesnil
— Propleural seta absent. Parafrontals not meeting, interfrontal area well developed.
Antennae not unusually small, antennal axis distinctly above mid-eye level. Mid
tibia with only one strong ad seta. [Parasites of Polistes] : 3 : . 2
2 Facial ridges strongly setulose for most of their height . ANACAMPTOMYIA Bischof
— Facial ridges bare (except for usual small setulae immediately above vibrissae)
EUVESPIVORA Baranov

Tribe STURMIINI

The Sturmiini are very well represented in Australia and Tasmania, the fauna
including many genera that are widely distributed elsewhere in the Old World
as well as several distinctive endemic genera. Several undescribed species are
known, and some of these may require additional new genera for their satisfactory
placement.

The group is moderately distinctive among the goniines with large pre-alar seta, and most

forms possess the following features in combination. Vibrissae inserted at least slightly
above (usually well above) the epistomal margin; gena wider than the profrons; 3 + 4 dc

92 KR. W. CROSSKEY

setae; humeral setae arranged as three in a straight line with one in addition set forwards of
the main line (this fourth seta occasionally missing); lower calypter with well defined inner
posterior angle and inner margin closely following edge of the scutellum. Other tribes possess
some of these characteristics, or some members of them do, but usually not all simultaneously
(for example, Winthemia and Carcelia have a sturmiine type of lower calypter in which the
inner edge abuts closely to the scutellum). All Australian Sturmiini, in addition to the
characters mentioned, have a strong submedian v seta on the mid tibia and have the ventral
surface of the second costal sector bare.

The endemic forms include a well-developed group in which the parafacials are
strongly haired, the eyes densely haired, the ocellar setae absent, the lateral
scutellar setae absent (exceptions in Polychaeta), and in which there is only one
definite pair of reclinate orbital setae. In some of these forms the antennae are
exceptionally long and the second aristal segment elongate, but such forms are
interconnected by intermediates with other forms in which the antennae are not
specially elongate and both basal aristal segments short. At present several
genera (e.g. Tvitaxys, Eurygastropsis, Polychaeta) are recognized in this complex,
but future study may well suggest that some of these genera ought to be
amalgamated. FEurygastropsis, for instance, is hardly distinguishable from
Tritaxys (syn. Goniophana) by more than the abdominal pollinosity and colour.
The genus Polychaeta exhibits some baffling variability in the scutellar setae;
normally in sturmiines these are extremely stable in strength and complement,
but in Polychaeta what appears to be the same species may have no lateral scutellar
setae at all, or may have one or two pairs extremely strongly developed (or even
be bilaterally asymmetrical in bristle complement in the same specimen).

The non-endemic forms found in Australia include well known genera of the
palaeotropics such as Palexorista, Sisyropa, Sturmia and Zygobothria. Genera
such as these occur (as would be expected) mainly in Queensland, though some
are represented widely throughout Australia; as a rule the non-endemic sturmiine
genera are unrepresented in Tasmania, though Blepharipa at least occurs there.
The widespread genera Drino and Pales are found in New Guinea, and their eventual
discovery in Queensland seems a possibility; they have accordingly been entered
in the key that follows.

The genus Winthellia (described by Mesnil under the preoccupied name T/yellina)
is here placed in the Sturmiini, although described as a _ winthemiine. Its
characters appear to ally it closely with Tasmaniomyia, which has all the main
diagnostic features of Sturmiini.

Key to AUSTRALIAN GENERA OF STURMIINI

1 Wing cell R, normal, distinctly open at the wing margin . : - ‘ : 2
Wing cell RF; closed just at or before the wing margin ; 24

2 Wings cisenieuy bicolorous, creamy yellowish near base and suffused with brown

anteromedially. Interfrontal area very reduced, under half as wide at mid point

as parafrontal. Apical scutellar setae exceptionally strong and horizontal, as
large as or larger than the subapical scutellar setae : : PALIANA Curran

— Wing not so, hyaline or at most only faintly yellowish or brownish anterobasally.

Interfrontal area not exceptionally narrow, subequal in width to parafrontal or

at least half as wide. Apical scutellar setae usually weak or very weak (some-

|

|

TACHINIDAE OF AUSTRALIA 93

times absent) and directed at least slightly upwards, if strong and horizontal

then weaker than subapical scutellar setae. ‘ é . : ‘ 3
Eyes haired (hairing long, dense and very conspicuous) : : 4
Eyes bare (a few very minute hairs sometimes visible under high Raamnertien : 13

Head without definite reclinate orbital setae. Frontal region bulbously convex
and with very reduced frontal setae (fig. 21 in Malloch, 1929b). Head ground
colour bright orange-yellow with blackish ocellar triangle. Legs unicolorous
orange-yellow. Abdomen thickly greyish white pollinose with a pattern of
black spots. Each side of venter of abdominal tergites 3 and 4 in g with a
dense hair fascicle - : ARRHENOMYZA Malloch
Head with one or more pairs of strong reclinate orbital setae. Frontal region not
bulbously convex (distinctly flattened when seen in profile) and with well developed
frontal setae. Head ground colour not so. Legs black or brownish black
(except in Zebromyia). Abdomen without such pattern. g without dense hair
fascicles on abdominal venter - 5
Facial ridges very wide and covered with several irregular rows of small strong
setulae (setulae only slightly increasing in size towards front edge of ridge).
Parafacials haired at upper ends immediately below lowest frontal setae.
Prescutum and scutum each with a transverse row of four small black spots
anteriorly (distinct against the yellowish grey pollinose background). Abdomen
conspicuously patterned, T3—-T5 greyish yellow pollinose with dark marks in
form of a pair of wide blackish or dark reddish brown triangles on T3 and T4
and a pair of small round black spots at tip of T5. Head with reddish ground
colour and chequered silvery white aan that appears to shift with direction
of light. Legs reddish yellow : : ZEBROMYIA Malloch
Facial ridges bare or setose; if setose then with ; a row ‘of strong setae on front edge
and only one or two inconspicuous setulae outside of the main row. Parafacials
either completely haired or completely bare. Prescutum and scutum without
such pattern. Abdomen without such pattern. Head not so. Legs black
or brownish black : - ‘ : : : : 6
Parafacials completely haired. Ocellar setae absent. Always fewer than four
sternopleuralsetae. Lateral scutellar setae absent (except in occasional specimens
of Polychaeta) : : 4 é : : 3 i ; : :
Parafacials bare. Ocellar setae present or absent. Sometimes four s¢p/ setae.
Lateral scutellar setae present ; 9
Dorsum of abdomen shining black or eee dane red-brown eecepe for last cavibl
tergite (T5) which is thickly coated with uniform whitish pollinosity (pale last
abdominal segment therefore strikingly contrasting with the rest of the abdomen)
EURYGASTROPSIS Townsend
Abdomen not so, ground colour reddish to blackish with thin whitish pollinosity
(sometimes appearing patchily distributed) on all tergites, no segments shining
and last segment not contrasted in colour with remainder of abdomen. : 8
Arista thickened on more than its basal half, sometimes on most of its length:
second aristal segment normally at least 2-5 times as long as broad. Pollinosity
of abdomen tending, to naked eye, to appear more conspicuous anteriorly on
segments than elsewhere or to appear rather chequered. Smaller forms, length
8-IImm . ‘ TRITAXYS Macquart
Arista only thickened on ts basal half or less: second aristal segment not more
than twice as long as broad. Pollinosity of abdomen tending, to naked eye,
to appear as a very thin but evenly distributed bloom. Larger forms, length
I2-16mm . POLYCHAETA Macquart
[This genus is only — doubttally deans on Tritaxys]
Four sternopleural setae (2 + 2). Second aristal segment distinctly elongate
(about three times as long as broad or more) : : = : : : 10

“I

94

Io

II

EZ

13

14

T5

16

17

18

R. W. CROSSKEY

Two or three sternopleural setae (I + 1 or 2 + 1). Second aristal segment not

atallelongate . : : c : : 3 : : : : II
One pair of reclinate orbital setae. Ocellar setae absent. Mid tibia with at least

two ad setae é . SISYROPA Brauer & Bergenstamm [part: S. faylori Curran]
Two or three pairs of reclinate orbital setae. Ocellar setae present, strong. Mid

tibia with one ad seta . : PARADRINO Mesnil
Facial ridges strongly setose 4 miOEE of (hem: height. One pair of reclinate orbital

setae. Dorsum of thorax shining bluish black, without pattern ; 12

Facial ridges bare. Two pairs of reclinate orbital setae. Dorsum of aire
yellowish grey to golden pollinose with a bold pattern of four black vittae on
prescutum, and a large black anteromedian area and pair of black sublateral
vittae on the scutum . : CALOZENILLIA Townsend

Ocellar setae absent. One strong ole ad seta on mid tibia . PALIA Curran

Ocellar setae present. Two or more strong ad setae on mid tibia

PALES Robineau-Desvoidy

Parafacials completely covered with long strong hair (hairing very conspicuous).
Ocellar setae directed outwards and not at all forwards. Lateral scutellar setae
absent. Parafrontals covered with long strong inclinate setulae that are nearly as
strong as the frontal setae. dG with very strong external vertical setae

PARAGONIA Mesnil

Parafacials bare or with rather weak inconspicuous hairing confined to upper
halves. Ocellar setae proclinate or absent (slightly divaricate in Wznthellia).
Lateral scutellar setae present or absent. Parafrontals usually without such
strong setae outside the frontal setae. ¢@ usually without definite external
vertical setae : 14

Lateral scutellar setae aliseae Geneelinen therefore ee only tree nae of mare
setae, the basals, subapicals and apicals). Upper halves of parafacials at least
partially haired. Three sternopleural setae. ¢G abdomen without secondary
sexual modified hair . ‘ 15

Lateral scutellar setae present (scutellum with a total BE Pate or ive aos of marsuel
setae). Upper halves of parafacials with or without hairing. Three or four
sternopleural setae. ¢G abdomen usually with some or most of the hair of the
venter of T4 (sometimes also T3) modified . : : : : . : 17

Ocellar setae present. Facial ridges bare (except for the usual few setulae on
lowermost quarter immediately above vibrissae). Abdominal tergites 1 + 2
and 3 each with a pair of strong median marginal setae. One or both of inter-
mediate abdominal tergites with discal setae. Upper ae with irregular
black setulae behind the postocular row : 16

Ocellar setae absent. Facial ridges strongly Senos io more Pie half fee
height. Abdominal tergites 1+ 2 and 3 without median marginal setae.
Intermediate abdominal tergites without discal setae. Upper aeae without
black setulae behind the postocular row ANAMASTAX Brauer & Bergenstamm

Mesonotum shining bluish black. Abdomen shining blackish green with traces of
reddish ground colour (not noticeably pollinose) TASMANIOMYIA Townsend

Mesonotum greyish white pollinose with four narrow blackish vittae. Abdomen
blackish with chequered whitish pollinosity conspicuous on intermediate tergites

WINTHELLIA Crosskey
[This genus and Tasmaniomyia are not separable on any convincing characters
and may need to be synonymized when better known]
Prosternum bare. Facial ridges with strong downcurved setae on more than

half their height : : BLEPHARELLA Macquart
Prosternum haired or seailess Baal Pies ae (except for usual very small
setulae immediately above vibrissae) . ‘ ‘ ‘ 18

Head with two pairs of reclinate orbital setae (upper one sometimes weak) ; : 19

——————— Cl Or rrr TOO Fr

19

20

21

22

23

TACHINIDAE OF AUSTRALIA 95

Head with one pair of reclinate orbital setae (strong and isolated, but 2 sometimes
with a smaller erect seta ventrad and mesad of the true reclinate orbital) . : 22
Gena conspicuously broad, two-fifths or more ofeye height. Facial ridges prominent
at least lower half well visible in profile. Parafacials about three times as wide
as third antennal segment. Three sternopleural setae (exceptionally four but
if so not arranged 2 + 2). Basal node of vein R,,,; with four or more small
setulae. Mid tibia with at least two strong ad setae. Head thickly golden
pollinose. Dorsum of thorax bright yellow or golden pollinose with four bold
black vittae (inner pair shorter than outer pair) : UGIMEIGENIA Townsend
Gena narrow, not exceeding one-sixth of eye height. Facial ridges not visible in
profile or only just visible immediately above vibrissae. Parafacials less than
twice as wide as third antennal segment. Four sternopleural setae (2 + 2).
Basal node of vein Ry, , with one setula (rarely also with one minute super-
numerary hair). Mid tibia with one ad seta (except in Zygobothvia atropivora).
Head not thickly golden te at most only partially pale yellow. Dorsum
of thorax not so - 20
Ocellar setae very strong, subequal i in size to reclinate orbital setae, inserted slightly
in front of anterior ocellus. Parafacials totally bare. Mid tibia with one or
two ad setae é : ZYGOBOTHRIA Mik
Ocellar setae absent or very weak and much smaller than reclinate orbital setae,
when present not inserted at all forwards of the anterior ocellus. Parafacials
finely haired on upper parts (bare in Dvino which is not yet positively known
from Australia). Mid tibia with one ad seta . : 3 : A : : 21
Parafacials finely haired on upper parts (Text-fig. 51), hair sometimes extending
slightly on to lower halves, occasionally only a few hairs immediately below
lowest frontal setae but parafacials never entirely bare. Ocellar setae present,
weak and wiry. Each side of venter of fourth abdominal tergite of ¢ with
well defined hair-patch, usually large and dense with hair very long and converging
into a distinct fascicle : : : : : . PALEXORISTA Townsend
Parafacials completely bare. Ocellar setae absent. Each side of venter of fourth
abdominal tergite of g with unmodified hair or with a large area of short fine
close-set hair not formed into a definite fascicle : . DRINO Robineau-Desvoidy
Scutellum with subapical setae inserted very far apart, distance between their
bases conspicuously greater than that between a subapical seta and its corres-
ponding basal seta. Apical scutellar setae very strong, horizontal or almost so,
nearly as strong as lateral setae. Four sternopleural setae (2 + 2). Abdomen
of 3 with a large area of very long dense hair around the sides of T4
STURMIA Robineau-Desvoidy
Scutellum with subapical setae not very widely separated, distance between their
bases less than or at most subequal to that between a subapical seta and its
corresponding basal seta. Apical scutellar setae very weak and directed at least
slightly upwards, very much weaker than lateral setae. Three or four sterno-
pleural setae. Abdomen of $ with or without such hair : 23
Abdomen with last tergite conspicuously small in relation to preceding tergite
(T5 usually not more than about half as long as T4). Three or four sternopleural
setae (normally three in g and four in 9, but males with four and females with
three occur). Lateral scutellar setae very strong, subequal in size to basals
and subapicals (often a second pair of laterals developed in addition to main
pair). Abdomen of ¢ with large area of very dense long hair around sides of
T4. Txr+2 and T3 always without median marginal setae. Larger forms
(length 9-15 mm) : : : BLEPHARIPA Rondani
Abdomen with last tergite Hoe gue small in relation to preceding tergite
(T5 subequal in length to T4 in ¢, about two-thirds as long in 9). Four sterno-
pleural setae in both sexes. Lateral scutellar setae well developed but distinctly

96 R. W. CROSSKEY

weaker than basals or subapicals (never a second pair of laterals). Abdomen
of f with hair on T4 unmodified. Tr + 2 and T3 with or without median marginal
setae. Smaller forms (length 6-8 mm)
SISYROPA Brauer & Bergenstamm [part: undetermined spp.]
24 Eyes bare. Abdomen patterned, T3—T5 thickly coated with whitish pollinosity
except for a shining black transverse fascia on each intermediate tergite which
is narrowly broken in the mid line but on each side extends round on to the
ventral surface of the tergite. Legs orange-yellow with blackish brown tarsi
QUADRA Malloch
— Eyes haired. Abdomen without such pattern, appearing generally blackish or
reddish black with an inconspicuous coating of yellowish grey pollinosity.
Legs mainly black or brownish black, only tibiae reddish yellow
‘Quadra’ dissimilis Malloch (? gen. n.)

Tribe GONIINI

This small tribe is easily distinguishable from all other members of the subfamily
Goniinae occurring in Australia by the strong reclinate ocellar setae (in other
goniine tribes the ocellars are either proclinate or absent). It is represented in
eastern and western Australia but appears to be absent from Tasmania. Two
genera are known from Australia, both of which have an extensive distribution
in Africa and the Oriental Region; one of them (Pseudogonia) occurs also in the
southern Palaearctic and eastwards as far as New Guinea and the Solomons.

KrEy TO AUSTRALIAN GENERA OF GONIINI

1 Three sternopleural setae (2 + I, one of anterior pair sometimes almost hair-like).
Scutellum without lateral setae (therefore with only two pairs of strong horizontal
marginal setae). Parafacials bare. Facial ridges setulose on most of their height

GONIOPHTHALMUS Villeneuve

— Four sternopleural setae. Scutellum with a pair of extremely strong lateral setae
(therefore with three pairs of horizontal marginals). Parafacials strongly but
sparsely haired (hairing nearest facial ridges setiform). Facial ridges bare

PSEUDOGONIA Brauer & Bergenstamm

Tribe ERYCIINI

This tribe is little more than a useful aggregate of genera to recognize
temporarily, until more is known of the biology of the included forms (especially
their reproductive habit). Almost certainly the tribe is polyphyletic and will
later need to be dismembered and its constituent genera reassigned to freshly
defined tribes. At the moment the tribe contains those genera of Goniinae with
large pre-alar seta that will not satisfactorily fit into other tribes (at least, not
without widening the definitions of these and destroying the homogeneity they
have at present). In practice if a goniine with large pre-alar setae does not show
the recognizable facies of a winthemuine, carceliine or sturmiine then it almost
certainly belongs in this tribe and its genus is most likely to be recognizable from
the eryciine generic key.

TACHINIDAE OF AUSTRALIA 97

The key to Australian genera here given is, however, only a preliminary one.
The Australian fauna contains many undescribed forms that will have to be fitted
into the Eryciini when they are named. Eryciine forms are rather well developed
in Australia, and most of the genera are endemic; some essentially extra-limital
genera (such as Aplomya and Erythrocera) occur, and it is likely that other
Oriento-Papuan genera (such as Aneogmena) will eventually be found in Queensland.
Some of the Australian eryciines, like Awstronilea, are extraordinarily similar to
Palaearctic counterparts, and the Tasmanian fauna includes several undescribed
forms that belong in the subgroup Baumhaueriina (a segregate occurring elsewhere
in the Palaearctic Region and southern Africa); the Tasmanian forms referred to
are recognized by the possession in the females of a pair of outwardly-directed
prevertical setae, and are the only Australian Goniinae known that possess
prevertical setae (except for the species of Spoggosia).

It may be useful to note that all the Australian eryciines so far known have the
lower surface of the second costal sector bare, and all have at least three setae on
the humeral callus, these standing more or less in a straight line (there are no
Australian eryciines, so far known at least, that have three humeral setae set in a
well defined triangle: three humerals in a triangle occur in Walker’s type of
Tachina calliphon, which is an eryciine, but there is uncertainty as to whether
this specimen is genuinely Australian).

The genus Bactromyiella is very difficult to place satisfactorily but is here
retained in the Eryciini; the small pra seta suggests that it ought possibly to be
placed in the Blondeliini, and the general facies and haired barette suggest that it
would not be inappropriate to treat it as a winthemiine.

The eryciine genera Aneogmena and Dolichocolon are included in the following
key as they occur in New Guinea, and may possibly occur in Australia.

Kry To AUSTRALIAN GENERA OF ERYCIINI

1 Wing with cell R; closed well before the wing margin and long-petiolate. Last
section of vein Cu, conspicuously longer than cross-vein m—cu. Two presutural
dorsocentral (prst dc) setae (very weak third setula sometimes developed between
two main setae). Apical scutellar setae directed almost straight upwards

PHOROCEROSTOMA Malloch

-— Wing with cell R; open at wing margin or just closed at the margin (not at all
petiolate). Last section of vein Cu, shorter than, or subequal in length to,
m-—cu. Three presutural dorsocentral setae (except in Aneogmena and Evythrocera
in which normally two). Apical setae of scutellum horizontal or directed
obliquely upwards, sometimes absent. . . : ; : : : : 2

2 Mid tibia without a submedian v seta. Barette haired along its length (sparsely
in 9). Sexes strongly dimorphic in colour and pattern: ¢ with thorax and first
abdominal segment black and remainder of abdomen bright golden orange or
orange-red; Q black with pleural regions, two fasciae on mesonotum, apex of
scutellum, and transverse band on each abdominal tergite 3-5, greyish yellow
to bright golden ; : BACTROMYIELLA Mesnil

-— Mid tibia with a submedian v ‘seta. Barette bare or with only one or very few
hairs anteriorly (more extensively haired in Tevetvophova). Sexes not strongly
dimorphic in colour or pattern, never as described above . : : ‘ 3

G

98

Io

R. W. CROSSKEY

Mesonotum and abdomen with a bold pattern of small black spots against a very

pale greyish yellow pollinose background, pattern comprised as follows: prescutum

with pair of rounded sublateral spots and anteromedian black mark, scutum

with prescutellar black spot; abdomen with Tr + 2 black and a median and a

pair of sublateral black spots on T3 and T4. Legs uniformly reddish yellow.

Presutural intra-alar seta absent, two or three postsutural intra-alar setae, ia

setae therefore o + 2-3. Head profile as Text-fig.50 . METAPHRYNO Crosskey
Nondescript forms without such pattern. Legs black or partly so, if mainly

reddish yellow then at least the tarsi black-brown. Intra-alar setae always

1 + 3. Head profile different : . 4
Wings distinctly bicolorous, creamy yellowish a near base aad Sateed with ieows

anteromedially. Interfrontal area very reduced, not more than half as wide as

a parafrontal. Apical scutellar setae exceptionally strong, as large as or larger

than the subapicals; subapical scutellar setae widely separated, bases much

wider apart than distance from insertion of a subapical seta to its corresponding

basal seta : : (PALIANA Curran, tribe Sturmiini]
Wings more or less completely hyaline, at most slightly brownish anterobasally.

Interfrontal area not conspicuously narrow, as wide as or only slightly narrower

than a parafrontal. Apical scutellar setae (when present) much weaker than

subapicals; subapical scutellar setae not unusually widely separated, distance

between bases at most only a little greater than distance from base of one subapical

to its corresponding basal seta : 5
Three post dc setae. Basal node of Ry,; fis a Tow Poe BL 7 demic: that extends

nearly half way to cross-vein y-m. All frontal setae of J very strongly reclinate.

Rather elongate black and silvery forms (with characteristic appearance of

scutellum which is sharply divided into silver-and-black halves which reverse

in appearance with direction of light) . ANEOGMENA Brauer & Bergenstamm
Four post dc setae. Basal node of R,,; with only 1-3 setulae. Frontal setae all

or mainly inclinate. Notsuch forms . : : : : ; : ‘ 6
Four sternopleural setae . : : : : ; - - : . a
Two or three sternopleural setae : - ‘ : : : : : 8

Mid tibia with one strong isolated ad seta. Eyes densely haired. Abdominal T5
unusually small, less than half as long as T4. Scutellum with apical setae
stronger than laterals, crossed and horizontal . APLOMYA Robineau-Desvoidy

Mid tibia with two strong ad setae. Eyes virtually bare. Abdominal T5 normal,
nearly as long as T4. Scutellum with apical setae weaker than laterals and
directed obliquely upwards . : ‘Bactromyia’ crassiseta Baranov (? genus)

Parafacials covered with very long conspicuous hair: haired area extending to
ventral ends of parafacials and more or less continuous with hairing of the genae.

Two sternopleural setae. Q with only one pair of proclinate orbital setae
PSEUDALSOMYIA Mesnil

Parafacials usually bare, if some hairing present then either confined to upper
halves of parafacials or very short and inconspicuous. Three sternopleural
setae (except in Palia and some specimens of Tevetvophora). Q with two pairs

of proclinate orbital setae. : 9
Mid tibia with one ad seta (very strong ands in pabmredices position) : 10
Mid tibia with two or more ad setae (setae additional to main one sometimes aeoe

small in g of Tevetvophora) . II

Intermediate abdominal tergites with discal setae. Ocellar setae present. ‘Upper
occiput with black setulae behind the postocular row. Hind tibia with a pd
preapical seta in addition to the normal d and ad preapicals. Eyes sparsely
short-haired. Femora reddish yellow (at most only slightly blackish near the
apices) : - . ERYTHROCERA Robineau-Desvoidy

Intermediate Abdomen es wana discal setae. Ocellar setae absent.

II

12

13

14

15

TACHINIDAE OF AUSTRALIA 99

Upper occiput without black setulae behind the postocular row. Hind tibia
without a pd preapical seta. Eyes densely long-haired. Femora black-brown
[PALIA Curran, tribe Sturmiini]
Parafacials finely and inconspicuously haired (especially on middle part of their
height). Hind tibia with a pd preapical seta in addition to the normal d and
ad preapicals. Antennae unicolorous bright orange. Legs reddish yellow
(except for the tarsi). Abdomen rather uniformly covered with golden pollinosity
in g and ashy grey pollinosity in 9. Lateral scutellar setae absent
CHLOROGASTROPSIS Townsend
Parafacials bare or with sparse stiff hairs on upper parts. Hind tibia without pd
preapical seta in addition to the d and ad preapicals. Lateral scutellar setae
present or absent. Antennae not unicolorous orange, at least the third segment
extensively brown. Legs largely blackish brown, the femora as well as the tarsi
at least partly dark (except in Aprotheca). Abdomen not so (but extensively
yellowish pollinose in A protheca 3) : 12
Scutellum without lateral setae. Apical scutellar setae well developed and crossed
(one of the pair absent in occasional ee Intermediate abdominal tergites
without discal setae . - - c : : : 2 13
Scutellum with a pair of long lateral setae. Apical scutellar setae absent or minute,
fine and divergent (? Apvotheca). Intermediate abdominal tergites with or
without discal setae. , - : - - - 14
Upper half of parafacials with ‘stiff setulose hairs. Facial ridges bare and widely
visible in profile. Eyes densely haired. Vibrissae well above level of epistomal
margin. Upper occiput with an irregular row of black setulae behind the
postocular row . AUSTROPHRYNO Townsend
Parafacials completely bare. Facial ridges with very strong downcurved setae
on nearly their entire height. Eyes almost bare (some very short sparse hairs
visible under high power magnification). Vibrissae level with epistomal margin.
Upper occiput without black setulae behind postocular row
DOLICHOCOLON Brauer & Bergenstamm
Legs black or brownish black. Facial region not noticeably warped forwards at
the epistome. Facial ridges with strong downcurved setae up most of their
height < : ; : : : : : : 15
Legs reddish yellow (except for dark tarsi). Facial region strongly warped forwards
at the epistome. Facial ridges bare (except for the usual few setulae immediately
above vibrissae) . APROTHECA Macquart*
Head in profile very strongly sabitniccoeilan (Text-fig. a: Intermediate abdominal
tergites each with a pair of strong erect discal setae. Upper occiput with black
setulae behind the postocular row. Abdominal T5 of 2 normal. Barette bare
AUSTRONILEA Crosskey
Head in profile not strikingly triangular, only a little shorter at the vibrissal axis
than at the antennal axis. Intermediate abdominal tergites without discal
setae, or one very feeble pair of discals haphazardly developed on T4. Upper
occiput without black setulae behind the postocular row. Abdominal T5 of 9
of extraordinary elongate conical shape (Text-fig. 95), preceding two tergites
correspondingly very short and relatively wide. Barette haired
TERETROPHORA Macquart

* The genus Aprotheca is known only from Macquart’s types that are in very bad condition. They
appear to have only three postsutural dorsocentral pores and the de complement in Aprotheca may
be 3 + 3.

100 Rk. W. CROSSKEY
PART II—A TAXONOMIC CATALOGUE OF THE AUSTRALIAN TACHINIDAE

INTRODUCTION

The only pre-existing published catalogue of the Australian Tachinidae is that
of Malloch (19280), in which he listed eighty-eight genera and two hundred and
twenty-eight species. Malloch’s list was essentially only a very preliminary one
derived solely from the literature, and Malloch was well aware of its limitations,
as is evident from the following paragraph in the preamble to his catalogue which
deserves quotation: ‘It will be seen that there is quite an array of species included,
but it is not to be assumed that all names standing in the list as apparently valid
species are in that category. I have not attempted to synonymize species, except
where previous workers have already done so, but I know many of the included
names will have to be sunk as synonyms of previously described forms appearing in
the catalogue. The work of weeding out such synonyms can only be done by someone
who will undertake a comprehensive study of the family, and definite identifications
will usually depend upon an examination of the type specimens of the species’.

In the forty-four years that have elapsed since Malloch’s catalogue there has
been considerable, though spasmodic, interest in the taxonomy of Australian
Tachinids, and many Australian genera and species have been described since
that time (many of them by Malloch himself in his post-1928 papers); in addition,
many non-endemic genera have been recognized in the Australian fauna. In
the new catalogue that follows a total of 136 genera are treated as valid, and
421 species are listed as valid on the evidence available at the moment (there are,
of course, many undescribed genera and species known in museum collections and
the figures given are for named taxa only). A large number of names, both
generic and specific, are treated as synonyms.

The catalogue is based on a study of very nearly all of the types of Australian
Tachinidae that are still in existence (the types are lost or missing in the cases
of only 29 out of a total of 487 nominal species-group taxa with an Australian
provenance). These studies of the types (including those of the type-species of
genera) have made it possible to weed out many of the obvious synonyms,
particularly among generic names but very often in the case of specific names
also. They have also made it possible to make reliable assignments of described
species to currently recognized genera, many of the placements so made involving
new combinations (as listed on p. 158). It cannot be claimed that all cases of
synonymy between specific names have been unmasked, and it is likely that future
critical generic revisionary work will reveal some additional specific synonyms.
(In difficult genera where specific criteria are uncertain at present it has been
considered best to maintain some or all of the specific names as valid until their
true status can be elucidated: cases of suspected synonymy have, however, been
indicated.)

EXPLANATORY INFORMATION ON THE CATALOGUE FORMAT

ARRANGEMENT OF TAXA AND NAMES. Subfamilies are placed in the conventional
order beginning with Phasiinae and ending with Goniinae. Tribes are in

TACHINIDAE OF AUSTRALIA 101

systematic order within subfamilies (rather arbitrary positional placements being
made for the more aberrant tribes). Genera, subgenera and species treated as
valid are listed alphabetically within their higher category, as there is no accepted
systematic order for their placement. Nomenclatorial synonyms are listed
chronologically under their respective valid names. Incorrect subsequent spellings
and misidentifications are listed after the nomenclatorial synonyms (if any), and
each entry of a misidentification is placed in square brackets.

CITATION OF NAMES AND REFERENCES. In each entry the name (generic,
subgeneric or specific) is cited first, followed by its author and date of publication
and the page reference to the work in which the name appeared; the author, year
date (letter-suffixed if more than one work by the same author in any year), and
page are always sufficient for obtaining the complete reference to the work containing
the name from the bibliography. Where a work is best known from a separately
paginated reprint version the reprint page reference is given in parentheses
immediately after the journal page reference.

The name of the original genus to which any species-group name was assigned
when first published is shown in parentheses immediately after the page reference,
but only when different from the present generic assignment (if no generic name
follows the page reference it is always to be inferred that the species-group taxon
in question was originally placed in the same genus as that in which it is here
listed). It is always clear from this convention whether or not any species remains
in its original genus, and parentheses have intentionally not been placed around
the authors’ names for those species that are no longer in their original genera
(mainly because the custom of bracketing around the names of authors of transferred
species does not lend itself to this type of catalogue in which both year date and
page reference are given).

Spellings of names accord with the rigid requirements of the International Code
of Zoological Nomenclature. Ordinarily the original spelling is the correct one
for each name, but as a few of the Australian tachinid genera have a neuter gender
(e.g. the names ending in -stoma and -soma) it has sometimes been necessary to
change the endings of adjectival specific names associated with such generic
names (to accord with Article 30 of the Code). For example, most species here
placed in Senostoma were originally described in genera with feminine gender
and the names originally had feminine endings; on being brought into combination
with Senostoma (neuter) the endings have had to be changed appropriately.
(Here it may be noted that I have no personal sympathy with Article 30, which
imposes an unnecessary burden on zoological nomenclature.) In some instances
a specific name when first published was associated with an incorrect subsequent
spelling of a generic name: when it appears helpful to do so, the incorrect spelling
is cited in parentheses after the page reference even though the species in question
remains in the same genus.

Citations of the mode of fixation of type-species accord with the Code, and are
‘original designation’, ‘monotypy’, and ‘subsequent designation’ (with a reference
to the designator given). Many monotypic genera when originally erected had

102 R. W. CROSSKEY

the type-species originally designated, in which case the mode of fixation is cited
solely as ‘original designation’, but the words ‘and monotypy’ are added in the few
special ‘gen. n., sp. n.’ situations of the kind covered by Article 68 (a) (i) of the Code.

Specific synonyms are indented but their citations and references are dealt
with in the same manner as the valid names. All valid names are printed in
bold-face italic type and junior synonyms in non-bold italics; other invalid names
(incorrect subsequent spellings, nomina nuda, misidentifications, infrasubspecific
names, junior primary homonyms in specific names, and junior homonyms in
generic names) are also printed in non-bold italics.

When a name was originally proposed for a genus but is now employed in the
catalogue as the valid name for a subgenus the words ‘as genus’ are appended in
parentheses after the page reference. Similarly, if a species-group name is now
employed in a different status from the original then the original status is indicated
in parentheses after the page reference, e.g. ‘as var. of ruficornis’.

TYPE-INFORMATION. The following information is given for the primary type
(holotype, lectotype or neotype) of each available species-group name listed in
the catalogue: type-status; sex of type; type-locality; type-depository and location;
a statement in the form ‘[examined]’ to show when the primary type has been
seen personally.

The following points should be noted about the data on primary types.

(1) Type-status. The primary type is cited as holotype if it is clear from the
original publication that only one specimen was available at the time of description
(whether designated in some way as ‘type’ or not), or if only one specimen is known
to exist and there is no published evidence as to how many specimens were
present in the type-series; and, of course, whenever a single specimen was
designated as type by the original author from a series of specimens. If lectotypes
and neotypes have been previously designated references are given to the place
of designation. A few lectotypes are newly designated in this work, and each
of these is appended ‘by present designation’ in the body of the catalogue. The
expression ‘Type(s)’ is used for the few cases in which the type-material is lost
or missing and it is not known from the original or later publications how many
specimens formed the original series.

(2) Sex of type. When the actual sex of the primary type is the opposite of
that cited in the original description the true sex is given first and the cited sex
after it, in the following manner: ‘Holotype ¢ [not 9]’. Such cases are infrequent
but occur now and then because of mis-recognition of sex by the earlier authors
(e.g. Walker, who was frequently unable to sex Tachinidae correctly). No sex is
cited at all in the very few instances in which no type-material has been found
or seen and there is no evidence of sex derivable from the original description
(e.g. Linnaemyia nigripalpus Tryon).

(3) Type-locality. In the citations of type-localities the larger territorial units
are cited before smaller ones, with the main unit (the Australian state or territory,
or if extra-Australian the country or island) shown in capital letters. If the
true provenance of the type differs from that cited in the original description then

TACHINIDAE OF AUSTRALIA 103

the true provenance is cited first and is followed by appropriate annotation in
square brackets. If the published locality is suspect (e.g. because later collecting
has not yielded specimens from the same area) but cannot be shown to be wrong
then it is placed in inverted commas, e.g. ‘TASMANIA’. Minor discrepancies
between the data on a primary type and that shown in the original publication are
suitably indicated if their importance warrants it. When citing type-localities
that are known more precisely than usual the following abbreviations have been
used: ‘Is’ = island; ‘mls’ = miles; ‘mt’ or ‘mts’ = mountain(s). Points of the
compass are abbreviated as N., E., S. and W. and combined as necessary.

(4) Iype-depository and location. These are shown in parentheses immediately
after the type-locality, with the abbreviation for the depository museum given
first and followed by the city. Ifa primary type is lost or has not been located
this is stated in parentheses after the type-locality (sometimes with some amplifying
words if pertinent). The abbreviations used for the museum depositories are
as follows:

AM Australian Museum, Sydney

AMNH American Museum of Natural History, New York

ANIC Australian National Insect Collection, Canberra

BMNH _ British Museum (Natural History), London

CNC Canadian National Collection, Ottawa

DEI Deutsches Entomologisches Institut, Eberswalde

MHN Musée d’Histoire Naturelle, Lille

MNHN Muséum National d’Histoire Naturelle, Paris

MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin
MRAC Musée Royal de |’Afrique Centrale, Tervuren

MZ Museo Zoologico ‘La Specola’, Florence

NM Naturhistorisches Museum, Vienna

NMV National Museum of Victoria, Melbourne

NR Naturhistoriska Riksmuseum, Stockholm

NSWDA New South Wales Department of Agriculture, Rydalmere
OM Queensland Museum, Brisbane

RMNH ~ Rijksmuseum van Natuurlijke Historie, Leiden
SPHTM = School of Public Health and Tropical Medicine, Sydney
USNM United States National Museum, Washington D.C.
UZM Universitetets Zoologiske Museum, Copenhagen
WADA Western Australian Department of Agriculture, Perth
ZM Zodlogisch Museum, Amsterdam

(5) Localities of extra-Australian type-species. It may conveniently be noted
here that statements of the original provenance of the type-species of genus-group
names are given only for those names that are based on extra-Australian
type-species. For example Calozenillia is based on a type-species from Sumatra,
and the locality ‘Sumatra’ is therefore given in parentheses at the end of the
generic entry. Localities are not given in the generic synonymies for the names

104 R. W. CROSSKEY

based on Australian type-species, as the latter are listed with their type-localities
in the catalogue of species that accompanies each generic (or subgeneric) name.

GEOGRAPHICAL DISTRIBUTION. An attempt has been made to provide a brief
synopsis of the known distribution of each species listed as valid in the catalogue.
Geographical data is given to the level of state or territory for Australia itself,
and to the level of island(s), country or zoogeographical region, as appropriate,
for species that occur extra-limitally as well as in Australia. Australian
distribution is given first, with the states listed alphabetically, followed by
extra-limital distribution (if any); the extra-limital distribution is listed with the
areas closest to Australia (e.g. New Guinea) first and more distant areas afterwards.
If there is only doubtful evidence of the occurrence of a particular species in any
Australian state, or if a published record from a state appears suspect, then the
state(s) in question are listed at the end of the Australian distribution and are
followed by a query mark: for example, Blepharipa fulviventris is recorded as
from ‘N.S.W., QLD, TASM. (?)’, this format indicating that records from New
South Wales and Queensland are considered confirmed but from Tasmania
doubtful (if records from two or more states are doubtful then a query mark
follows the entry for each such state). The abbreviations used for listing Australian
states and territories are as follows:

A.C.T. Australian Capital Territory S.A. South Australia

N.S.W. New South Wales Tasm. Tasmania
NA. Northern Territory Vict. Victoria
OLD Queensland W.A. Western Australia

Lord Howe Island is listed separately from New South Wales and is not
abbreviated.

NOMENCLATURAL CHANGES. It has been necessary to establish in the catalogue
several new generic and specific synonymies, many new combinations for
generically re-assigned species, and a few new names for preoccupied homonyms:
a summary of these changes is provided on p. 157. The usual bold-face
abbreviations have been used to signify these changes, viz. Comb. n. (new
combination), Nom. n. (new name) and Syn. n. (new synonym). New combinations
are only marked as such when considered taxonomically valid; species-group
names that are assigned for the first time to a particular generic taxon are not
marked as new combinations if they are also junior synonyms (as in these instances
there are no valid new binomina in use). The abbreviation Comb. n. when given
is placed after the type-locality data (which concludes the essentially nomenclatorial
matter) and before the distributional data in the body of the catalogue, and each
new taxonomically valid binomen is set out formally in the summary of new
combinations (p. 158).

MISCELLANEOUS ANNOTATIONS. Whenever it is necessary or desirable to call
attention to some specially pertinent point concerning a species-group name
appearing in the catalogue (e.g. to elucidate points of homonymy, possible

TACHINIDAE OF AUSTRALIA 105

whereabouts of types, mis-citations of dates, etc.) the additional annotation is
given in a separate paragraph immediately following the main entry.

SYNOPSIS OF THE CATALOGUE ARRANGEMENT OF TRIBES AND GENERA

The following synopsis is given to show at a glance the arrangement of tribes
and genera in the body of the catalogue. The genera listed are those considered
valid at the present time. The affinities of several Australian genera are
particularly obscure, and some have had to be assigned to tribes rather arbitrarily;
likewise, the appropriate subfamily assignments of some tribes are doubtful. All
such cases of doubtfully assigned genera and tribes are indicated by an asterisk
(*) against the name; taxa so marked are in special need of study to ascertain
their relationships more clearly.

Subfamily PHASIINAE

Tribe TRICHOPODINI

Pentatomophaga de Meijere
Saralba Walker

Tribe PHASIINI
Alophora Robineau-Desvoidy
Besserioides Curran
Efftayloria Malloch

Tribe CYLINDROMYIINI
Cylindromyia Meigen
Gerocyptera Townsend

Tribe LEUCOSTOMATINI
Leucostoma Meigen

Tribe EUTHERINI
Euthera Loew

Subfamily PROSENINAE (DEXIINAE)

Tribe PROSENINI
Acucera Malloch
Anatropomyia Malloch
Geraldia Malloch
Heterometopia Macquart*
Hobartia Malloch
Macropodexia Townsend
Ola Paramonov
Platytainia Macquart
Prosena Le Peletier & Serville
Pyrosenina Malloch
Rutilotrixa Townsend
Senostoma Macquart
Trichostylum Macquart

Tribe RUTILIINI
Amphibolia Macquart
Chetogaster Macquart

106 R. W. CROSSKEY

Chrysopasta Brauer & Bergenstamm
Formosia Guérin-Méneville
Prodiaphania Townsend

Rutilia Robineau-Desvoidy

Subfamily TACHININAE (MACQUARTIINAE)

Tribe PALPOSTOMATINI*
A palpostoma Malloch
Eustacomyia Malloch
Palpostoma Robineau-Desvoidy

Tribe MYIOTRIXINI*
Myiotvixa Brauer & Bergenstamm

Tribe ORMIINI*
Therobia Brauer

Tribe GLAUROCARINI*
Doddiana Curran

Tribe CAMPYLOCHETINI
Elpe Robineau-Desvoidy

Tribe VORIINI
Hyleovus Aldrich
Hystricovoria Townsend
Voria Robineau-Desvoidy

Tribe THELAIRINI
Halydaia Egger
Thelaiva Robineau-Desvoidy

Tribe MINTHOINI
Minthoxia Mesnil
Sumpigaster Macquart

Tribe NEMORAEINI
Nemoraea Robineau-Desvoidy

Tribe LESKIINI
Apatemyia Macquart
Demoticoides Mesnil
Exechopalpus Macquart
Rhinomyobia Brauer & Bergenstamm
Sipholeskia Townsend
Toxocnemis Macquart

Tribe ERNESTIINI
Amphitropesa Townsend*
Chlorotachina Townsend
Macrochloria Malloch
Neximyia Crosskey*

Tribe PARERIGONINI*
Austvalotachina Curran*
Leverella Baranov
Pygidimyia Crosskey
Zita Curran

eee

TACHINIDAE OF AUSTRALIA

Tribe LINNAEMYINI
Apalpus Malloch
Chaetophthalmus Brauer & Bergenstamm
Linnaemya Robineau-Desvoidy

Tribe TACHININI
Cuphocera Macquart
Eristaliomyia Townsend
Microtropesa Macquart

Subfamily GONIINAE

Tribe ACEMYINI
Ceracia Rondani

Tribe NEAERINI
Voriella Malloch

Tribe SIPHONINI (ACTIINI)
Actia Robineau-Desvoidy
Ceromya Robineau-Desvoidy
Peribaea Robineau-Desvoidy

Tribe BLONDELIINI
Anagonia Brauer & Bergenstamm
Compsilura Bouché
Deltomyza Malloch
Froggattimyia Townsend
Lecanipa Rondani
Lixophaga Townsend
Medinodexia Townsend
Monoleptophaga Baranov
Pareupogona Townsend
Paropsivora Malloch
Pilimyia Malloch
Trigonospila Pokorny
Zenargomyia Crosskey
Zosteromeigenia Townsend

Tribe EXORISTINI
Austrophorocera Townsend
Chaetoria Becker
Eozenillia Townsend
Exorista Meigen
Hillomyia Crosskey nom. n.
Spoggosia Rondani
Stomatomyia Brauer & Bergenstamm

Tribe ETHILLINI
Ethilla Robineau-Desvoidy
Mycteromyiella Mesnil*
Phorocerosoma Townsend

Tribe WINTHEMIINI
Crypsina Brauer & Bergenstamm
Nemorilla Rondani
Winthemia Robineau-Desvoidy

107

108 R. W. CROSSKEY

Tribe CARCELIINI

Argyrophylax Brauer & Bergenstamm
Argyrothelaiva Townsend

Carcelia Robineau-Desvoidy
Carcelimyia Mesnil

Tribe ANACAMPTOMYIINI

Anacamptomyia Bischof
Euvespivora Baranov
Koralliiomyia Mesnil*

Tribe STURMIINI

Anamastax Brauer & Bergenstamm
Arrhenomyza Malloch
Blepharella Macquart
Blepharipa Rondani
Calozenillia Townsend
Eurygastropsis Townsend
Palexorista Townsend

Pala Curran*

Paliana Curran*

Paradrino Mesnil

Paragonia Mesnil

Polychaeta Macquart
Quadra Malloch

Sisyropa Brauer & Bergenstamm
Sturmia Robineau-Desvoidy
Tasmaniomyia Townsend
Tritaxys Macquart
Ugimeigenia Townsend
Winthellia Crosskey
Zebromyia Malloch
Zygobothria Mik

Tribe GONIINI

Goniophthalmus Villeneuve
Pseudogonia Brauer & Bergenstamm

Tribe ERYCIINI

Aplomya Robineau-Desvoidy
A protheca Macquart
Austronilea Crosskey
Austrophryno Townsend
Bactromyiella Mesnil*
Chlorogastropsis Townsend
Evythrocera Robineau-Desvoidy
Metaphryno Crosskey
Phorocerostoma Malloch
Pseudalsomyia Mesnil
Tevetrophova Macquart

a __

TACHINIDAE OF AUSTRALIA 109

THE TAXONOMIC CATALOGUE

Family TACHINIDAE Robineau-Desvoidy
TACHINARIAE Robineau-Desvoidy, 1830 : 185. Type-genus: Tachina Meigen, 1803.

Subfamily PHASIINAE Robineau-Desvoidy
PHASIANEAE Robineau-Desvoidy, 1830 : 280. Type-genus: Phasia Latreille, 1804.

Tribe TRICHOPODINI Townsend
TRICHOPODINI Townsend, 1908 : 129. Type-genus: Tvichopoda Latreille, 1825.

Genus PENTATOMOPHAGA de Meijere

Pentatomophaga de Meijere, 1917 : 246. Type-species: Pentatomophaga bicincta de Meijere,
1917, by monotypy. (JAVA).

bicincta de Meijere, 1917 : 247. Holotype 9, Java (ZM, Amsterdam) [examined]. — QLp;
Java; NEw Britain.

Genus SARALBA Walker

Savalba Walker, 1865:114. Type-species: Savalba ocypteroides Walker, 1865, by monotypy.
(NEw GUINEA).

Pseudotrichopoda Malloch, 19334: 77. Type-species: Pseudotrichopoda varipes Malloch, 1933
[=Saralba ocypteroides Walker, 1865], by original designation.

bancrofti Paramonov, 1956 : 373. Holotype 3, QUEENSLAND, Eidsvold (ANIC, Canberra)
[examined].—Oxp. (Probably = ocypteroides).

ocypteroides Walker, 1865 : 114. Holotype g, NEw GuinEea (BMNH, London) [examined].—
QLD; NEw GUINEA.

varipes Malloch, 1933a : 78 (Pseudotrichopoda). Holotype g, QUEENSLAND, Cairns (DEI,
Eberswalde) [examined].

Tribe PHASIINI Robineau-Desvoidy
PHASIANEAE Robineau-Desvoidy, 1830 : 280. Type-genus: Phasia Latreille, 1804.

Genus ALOPHORA Robineau-Desvoidy

Alophora Robineau-Desvoidy, 1830 : 293. Type-species: Syvphus hemipterus Fabricius, 1794,
by subsequent designation of Coquillett (I910 : 505). (EUROPE).
Allophova. Incorrect subsequent spelling of Alophova Robineau-Desvoidy.

Subgenus ALOPHORELLA Townsend
Alophorella Townsend, 1912 : 45. Type-species: Theveva obesa Fabricius, 1798, by original
designation. (EUROPE).

Austrophasia Townsend, 1916f : 45. Type-species: Hyalomya rufiventyis Macquart, 1851, by
original designation.

IIo kK. W. CROSSKEW

aureiventris Curran, 19274: 165. Holotype 3, QUEENSLAND: Biloela (BMNH, London)
[examined]. — QLD.

chrysis Malloch, 1930a@:95 (Hyalomyia). Holotype g, WESTERN AUSTRALIA: Narragin
(SPHTM, Sydney) [examined]. Comb. n.—W.A.

costalis Malloch, 1929 : 284 (Hyalomyia). Holotype g, NEw SoutH WatLes: Woodford
(ANIC, Canberra) [examined]. Comb. n.-—N.S.W.

discalis Malloch, 1930a:95 (Hyalomyia). Holotype g, WESTERN AUSTRALIA: Geraldton
(SPHTM, Sydney). Comb. n.—W.A., QLD (?), Tasm. (?).

rufiventris Macquart, 1851 : 188 (215) (Hyalomya). Holotype g, ‘Tasmania’ (MNHN,
Paris) [examined].—Tasm. (?).

Subgenus HYALOMYA Robineau-Desvoidy

Hyalomya Robineau-Desvoidy, 1830: 298 (as genus). Type-species: Phasia semicinevea
Meigen, 1824 [=Phasia pusilla Meigen, 1824], by subsequent designation of Westwood
(1840 : 140). (EUROPE).

Hyalomyia. Incorrect subsequent spelling of Hyalomya Robineau-Desvoidy.

normalis Curran, 1927) : 355 (Strongylogaster). Holotype 9, QUEENSLAND: Palmerston
(DEI, Eberswalde) [examined]. Comb. n.-—QLp.

Subgenus MORMONOMYIA Brauer & Bergenstamm

Mormonomyia Brauer & Bergenstamm, 1891 : 388 (84) (asgenus). Type-species: Mormonomyia
laniveniris Brauer & Bergenstamm, 1891 [=Phasia argentifrons Walker, 1849], by subsequent
designation of Townsend (1916a : 8). (SouTH AFRICA).

basalis Malloch, 1930a : 96 (Hyalomyia). Holotype 3g, NEw SoutH Waters: Wahroonga
(SPHTM, Sydney) [examined]. Comb. n.-—N.S.W.

hyalis Malloch, 1930a : 96 (Hyalomyia). Holotype g, NEw SoutTH WaLEs: Como (USNM,
Washington) [examined]. Comb. n.—N.S.W.

lativentris Malloch, 1929a : 110 (Hyalomyia). Holotype g, QUEENSLAND: Cairns (USNM,
Washington) [examined]. Comb. n.— QLD.

lepidofera Malloch, 1929a: 111 (Hyalomyia). Holotype g, NEw SouTH WaLEs: Como
(USNM, Washington) [examined]. Comb. n.—N.S.W.

nigrihirta Malloch, 1929a: 112 (Hyalomyia). Holotype g, Victoria: Seaford (USNM,
Washington) [examined]. Comb. n.— QLD, Vict.

sensua Curran, 1927) : 354 (Strongylogaster). Holotype 9, QUEENSLAND: Palmerston (DEI,
Eberswalde) [examined]. Comb. n. — QLD.

Subgenerically unplaced species of Alophora s.1.

hippobosca Paramonov, 1958:594 (Hyalomyia). Holotype g, AUSTRALIAN CaPITAL
TERRITORY: Canberra (ANIC, Canberra) [examined]. Comb. n.—A.C.T., Vict.
This species is close to subgenus Alophorella but the parafrontals are bare outside the
frontal rows; it comes closest to the Palaearctic subgenus Phovantha Rondani.
nigrisquama Malloch, 1929a: 110 (Hyalomyia). Holotype g, NEw SoutH Wates: N.
Sydney, French’s Forest (AM, Sydney) [examined]. Comb. n.—N.S.W.
This species has the face and epistome flat in profile and the subgenus is uncertain.
It comes closest to the Palaearctic subgenus Brumpiallophora Dupuis.

Genus BESSERIOIDES Curran

Besserioides Curran, 1938 : 185. Type-species: Besserioides sexualis Curran, 1938
[=Catharosia varicoloy Curran, 1927], by original designation.

TACHINIDAE OF AUSTRALIA IIt

bancrofti Paramonov, 1958: 597. Holotype g, QUEENSLAND, Palm Is. (ANIC, Canberra)
[examined]. — QLD.

latifrons Paramonov, 1958 : 596. Holotype g, AUSTRALIAN CaPITAL TERRITORY, Bendora
(ANIC, Canberra) [examined]. — A.C.T.

varicolor Curran, 19274: 165 (Catharosia). Holotype g, QUEENSLAND, Bileola (BMNH,
London) [examined]. — QLD.

sexualis Curran, 1938: 185. Holotype g, QUEENSLAND, Laidley (BMNH, London)

[examined]. — Syn. n.

Genus EFFTAYLORIA Malloch

Taylovia Malloch, 1930a: 98. Type-species: Taylorvia testacea Malloch, 1930, by original
designation. [Junior homonym of Tayloria Bourguignat, 1889.]
Efftayloria Malloch, 1941 : 64. [Replacement name for Tayloria Malloch.]

testacea Malloch, 1930a : 98 (Tayloria). Holotype g, QUEENSLAND, Mt Molloy (SPHTM,
Sydney) [examined].—Q.ip. (Possibly = ochromyoides Walker, 1865, New Guinea).

Tribe CYLINDROMYIINI Townsend
CYLINDROMYIINI Townsend, 1912 : 48. Type-genus: Cylindromyia Meigen, 1803.

Genus CYLINDROMYIA Meigen

Cylindromyia Meigen, 1803 : 279. Type-species: Musca brassicavia Fabricius, 1775, by
monotypy. (EUROPE).

Ocypteropsis Townsend, 1916b : 630. Type-species: Ocyptera flavifyrons Macquart, 1851
[=Ocyptera bimacula Walker, 1849], by original designation.

[Ocyptera Latreille sensu authors (misidentification) |

angustissimifrons Paramonov, 1956 : 365. Holotype g, AUSTRALIAN CAPITAL TERRITORY:
Mt Gingera (ANIC, Canberra) [examined]. — A.C.T.
aterrima Paramonov, 1956 : 361. Holotype g, QUEENSLAND: Scotsville (ANIC, Canberra)
[examined]. — OLD.
atratula Malloch, 1930b : 314. Holotype 9, NEw SoutH WaLEs: Sydney (SPHTM, Sydney)
[examined]. — N.S.W.
bicoloy Bigot, 1885c:lv (Glossidionophora). Holotype 9, AustTratta (BMNH, London)
[examined]. — State unknown.
C. bicolor Bigot is a junior secondary homonym of C. bicolor Olivier, 1811. No new
name is proposed at the present time.
bimacula Walker, 1849 : 694 (Ocyptera). Holotype gj, Tasmania (BMNH, London) [examined].
—A.C.T., N.S.W., S.A., Tasm., W.A.
flavifrons Macquart, 1851 : 187 (214) (Ocyptera). Lectotype 3 (by designation of Crosskey,
I97I : 280), TasmANIA (MNHN, Paris) [examined]. Syn. n.
brunnea Malloch, 1930b : 315. Holotype g, WESTERN AUSTRALIA: Perth (ANIC, Canberra)
[examined]. — W.A.
dayi Paramonov, 1956 : 369. Holotype g, NEw Sourn Wates: Killara (ANIC, Canberra)
[examined]. — N.S.W.
fenestrata Paramonov, 1956 : 363. Holotype g, QUEENSLAND: Gayndah (AM, Sydney)
[examined]. — OLp.
hobartana Paramonov, 1956 : 368. Holotype g, Tasmania: Hobart (ANIC, Canberra)
[examined]. — Tas.
howeana Paramonov, 1956: 364. Holotype g, Lorp Hower Istanp (ANIC, Canberra)
[examined]. - Lorp Howe I.

112 Ri Wt GCROSSKEY

nigricosta Malloch, 1930) : 312. Holotype ¢ [two wings only remaining], NEw Soutu
WateEs: Kosciusko (SPHTM, Sydney) [examined]. — N.S.W.

rieki Paramonov, 1956 : 370. Holotype g, Tasmania: Eagle Hawk Neck (ANIC, Canberra)
[examined]. — Tasm.

rufifemur Paramonov, 1956 : 366. Holotype g, NEw SoutH WaLEs: Salisbury Downs-
Clifton Downs (ANIC, Canberra) [examined]. — N.S.W., OLp.

sydneyensis Malloch, 1930) : 314. Holotype2, NEw Soutu WaLEs: Sydney (ANIC, Canberra)
[examined]. —N.S.W.

tricolor Malloch, 1930b : 315. Holotype 9, New Soutu WateEs: Killara, Allowrie (SPHTM,
Sydney) [examined]. — N.S.W.

unguiculata Paramonov, 1956: 370. Holotype g, AUSTRALIAN CAPITAL TERRITORY:
Blundell’s (ANIC, Canberra) [examined]. — A.C.T., OLD., W.A.

westralica Paramonov, 1956: 367. Holotype g, WESTERN AUSTRALIA: King George’s
Sound (AM, Sydney) [examined]. — W.A.

Genus GEROCYPTERA Townsend

Gerocyptera Townsend, 1916e : 178. Type-species: Tvichoprosopa marginalis Walker, 1860, by
original designation. (MOLuccas).

tristis Bigot, 1878 : 45 (Ocypieva). Holotype g, ‘AusTRALIA’ (BMNH, London) [examined].
Comb. n. — AUSTRALIA (?).

Tribe LEUCOSTOMATINI Townsend
LEUCOSTOMINI Townsend, 1908 : 76. Type-genus: Leucostoma Meigen, 1803.

Genus LEUCOSTOMA Meigen

Leucostoma Meigen, 1803 : 279. Type-species: Ocypteva simplex Fallén, 1815, by subsequent
monotypy (Meigen, 1824 : 234). (EUROPE).

Aequia Malloch, 1930b : 325. Inadvertent error for Leucostoma, published in the statement
‘The preceding genus, Aeguwia, is distinguished from the present one [Hyalomyodes] by the
strong erect ocellar bristles, and the wing venation’. Here held to be a name first published
as a synonym and unavailable under Article 11(d) of the International Code of Zoological
Nomenclature. (See also Townsend, 1938 : 189).

simplex Fallén, 1815 : 240 (Ocyptera). Holotype 9, SwEDEN: Kalmar (NR, Stockholm)
[examined].—N.S.W.; widespread HoLarctic REGIONS, also NEOTROPICAL REGION.

Tribe EUTHERINI Townsend
EUTHERINI Townsend, 1912 : 49. Type-genus: Euthera Loew, 1866.

Genus EUTHERA Loew

Euthera Loew, 1866 : 46. Type-species: Eutheva tentatrix Loew, 1866, by monotypy. (UNITED
STATES OF AMERICA).
Macreuthera Bezzi, 1925 : 281 (as subg. of Euthera). Type-species: Euthera skusei Bezzi,

1925, by original designation.
rieki Paramonov, 1953 : 207. Holotype gf, QUEENSLAND: Warri Border Gate-Narylco (ANIC,
Canberra) [examined]. —- QLD.

TACHINIDAE OF AUSTRALIA 113

skusei Bezzi, 1925: 281. Holotype 9, QUEENSLAND: Eidsvold (whereabouts unknown,
possibly lost). — QLD.

Holotype should be in SPHTM, Sydney, but has not been located.

Unplaced species of Phasiinae

australasiae Malloch, 1930b : 325 (Hyalomyodes). Holotype 3, NEw SoutH WatEs: Sydney
(SPHTM, Sydney) [examined]. — N.S.W.

Placed by Malloch in New World genus Hyalomyodes Townsend but probably does not
belong there.

Subfamily PROSENINAE Townsend
PROSENINAE Townsend, 1892b : 273. Type-genus: Prosena Le Peletier & Serville, 1828.

Tribe PROSENINI Townsend
PROSENINAE Townsend, 1892b : 273. Type-genus: Prosena Le Peletier & Serville, 1828.

Genus ACUCERA Malloch
Acuceva Malloch, 1930b : 328. Type-species: Acucerva montana Malloch, 1930, by original
designation.

montana Malloch, 1930b : 328. Holotype gj, NEw Soutu WALEs: Blue Mts (SPHTM, Sydney)
[examined]. — N.S.W.

Genus ANATROPOMYIA Malloch

Anatropomyia Malloch, 1930a: 126. Type-species: Anatropomyia flavicornis Malloch, 1930,
by original designation.

flavicornis Malloch, 1930a : 127. Holotype §, NEw SoutH WatgEs: Ilford (SPHTM, Sydney)
[examined]. — N.S.W.

Genus GERALDIA Malloch

Gervaldia Malloch, 1930b : 327. Type-species: Gevaldia hirticeps Malloch, 1930, by original
designation.

hirticeps Malloch, 1930) : 328. Holotype g, WESTERN AuSTRALIA: Geraldton (SPHTM,
Sydney) [examined].-—S.A., W.A.

Genus HETEROMETOPIA Macquart

Heterometopia Macquart, 1846 : 298 (170). Type-species: Heterometopia argentea Macquart,
1846, by monotypy.
Cystometopia Townsend, 1926 : 531. Type-species: Heterometopia rufipalpis Macquart, 1847
[=Heterometopia argentea Macquart, 1846], by original designation.
H

114 R. W. CROSSKEY

argentea Macquart, 1846 : 298 (170). Lectotype ¢ (by designation of Crosskey, 1971 : 271),
Tasmania (MNHN, Paris) [examined]. — Tasm.
limbinevris [sic] Macquart, 1846 : 317 (189) (Omalogaster). Holotype 9 [not 9], TASMANIA
(MNHN, Paris) [examined]. Syn. n.
nitidus Macquart, 1846 : 318 (190) (Omalogaster). Type(s), TASMANIA (lost). Syn. n.
rufipalpis Macquart, 1847 : 90 (74). Holotype 3 [not 9], Austratia (BMNH, London)
[examined]. Syn. n.
analis Macquart, 1851 : 182 (209). Lectotype 2 (by designation of Crosskey, 1971 : 271),
TASMANIA (MNHN, Paris) [examined].
bella Paramonov, 1960 : 692. Holotype g, NEw Soutu WatEs: 1o mls SE of Braidwood

(ANIC, Canberra) [examined]. — A.C.T., N.S.W.
montana Paramonov, 1960 : 693. Holotype g, AUSTRALIAN CAPITAL TERRITORY: Bendora
(ANIC, Canberra) [examined]. — A.C.T., N.S.W., Vict.

Genus HOBARTIA Malloch

Hobartia Malloch, 1930a : 127. Type-species: Hobartia peculiaris Malloch, 1930, by original
designation.

peculiaris Malloch, 1930a:127. Holotype g, Tasmania: Hobart (ANIC, Canberra)
[examined]. — TAsM.

Genus MACROPODEXIA Townsend

Macropodexia Townsend, 1933 : 462. Type-species: Dexia longipes Macquart, 1846, by original
designation.

longipes Macquart, 1846 : 315 (187) (Dexia). Lectotype ¢ (by designation of Crosskey,
1971 : 266), TASMANIA (MNHN, Paris) [examined]. — N.S.W., Tasm., Vict.

Genus OLA Paramonov

Ola Paramonov, 1968 : 377. Type-species: Rutilia nigrithorax Macquart, 1851, by original
designation.
erasmusi Paramonov, 1968 : 381. Holotype g, Victoria: Warburton (NMV, Melbourne). —

Vicr.
insectaria Paramonov, 1968 : 380. Holotype g, AUSTRALIAN CAPITAL TERRITORY: Canberra

(ANIC, Canberra). —- A.C.T., N.S.W.
nigrithorax Macquart, 1851 : 190 (217) (Rutila). Lectotype ¢ (by designation of Crosskey,
1971 : 288), AUSTRALIA [publ. as ‘Oceania’] (MNHN, Paris) [examined]. — A.C.T., N.S.W.,

Tasm., VICT.
wilsoni Paramonov, 1954 : 281 (Chaetogaster). Holotype g, NEw SoutH Wa Es: Blackheath

(NMV, Melbourne). — N.S.W., Vict.

Genus PLATYTAINIA Macquart

Platytainia Macquart, 1851 : 178 (205). Type-species: Platytainia maculata Macquart, 1851,
by monotypy.

maculata Macquart, 1851 : 179 (206). Holotype 9, ‘Tasmania’ (MNHN, Paris) [examined]. —
N.S.W., Tasm. (?).

|
|

TACHINIDAE OF AUSTRALIA 115

Genus PROSENA Le Peletier & Serville

Calivrhoe Meigen, 1800 : 39. [Name suppressed by I.C.Z.N. Opinion 678 (1963 : 339)]
Prosena Le Peletier & Serville, 1828 : 499, 500. Type-species: Stomoxys siberita Fabricius,
1775, by original designation. (DENMARK).

arcuata Malloch, 1932a: 129. Holotype g, NEw Soutu Wa tEs: Killara, Allowrie (SPHTM
Sydney) [examined]. — N.S.W.
argentata Curran, 1927) : 348. Holotype g, QUEENSLAND: Kuranda (DEI, Eberswalde). —
QLD.
P. argentata Curran is a junior primary homonym of P. argentata Walker, 1859. No
new name is proposed at the present time.
bella Curran, 1927) : 349. Holotype g, QUEENSLAND: Cairns (DEI, Eberswalde).— QLp.
bisetosa Malloch, 1932a : 130. Holotype J, QUEENSLAND: Cairns district (SPHTM, Sydney)
[examined]. — QuD., N.S.W. (?).
conica Guérin-Méneville, 1831 : Plate 21, fig. 7, 7A; 1838: 298. Holotype gj, NEw Soutu
WateEs: Port Jackson (=Sydney) (MNHN, Paris) [examined]. —N.S.W., Orb.
albifrons Malloch, 1932a : 132. Holotype g, QUEENSLAND: Eidsvold (not located). Syn. n.
{argentata Curran sensu Malloch, 1930 (misidentification) ]
dimidiata Curran, 1938 : 188. Holotype 9, QuEENSLAND: Kuranda (SPHTM, Sydney)
[examined]. — OLD.
dispar Macquart, 1851 : 203 (230). Lectotype 9 (by designation of Crosskey, 1971 : 284),
‘Tasmania’ (MNHN, Paris) [examined]. — N.S.W., Taso. (?).
doddi Curran, 1927) : 347. Holotype g, QUEENSLAND: Herberton (DEI, Eberswalde).—N.T.,
Otp. (=siberita Fabricius according to Malloch, 1932a : 131).
dorsalis Macquart, 1847 : 97 (81). Holotype g, Tasmania (BMNH, London) [examined]. —
N.S.W., TasM.
parva Malloch, 1930a: 115. Holotype 9, New SoutH WatEs: Sydney (SPHTM, Sydney)
[examined]. Syn. n.
jactans Walker, 1858: 210 (Pachymyia). Holotype 2 [? AustTratia] (BMNH, London)
[examined]. —N.S.W., QLD.
macropus Thomson, 1869 : 531. Holotype g, NEw SoutH WaLEs: Sydney (NR, Stockholm)
[examined]. — N.S.W., Tas.
indecisa Malloch, 1930a: 116. Holotype g, NEw SoutH Wa Es: Barrington Tops (not
located). Syn. n.
Paratypes undoubtedly conspecific with missing holotype are in USNM, Washington,
and SPHTM, Sydney [examined].
malayana Townsend, 1926a: 25 (Calirrhoe). Lectotype g (by designation of Crosskey,
1969 : 91), SUMATRA: Fort de Kock (ZM, Amsterdam) [examined].—QtLp.; widespread
S.E. Asta. (Possibly = siberita).
P. malayana was considered by Curran (1938 : 189) to be a senior synonym of P. doddi.
Evidence is inconclusive.
marginalis Curran, 1938: 189. Holotype g, QUEENSLAND: Brisbane (SPHTM, Sydney)
[examined]. — QLD.
nigripes Curran, 1927) : 347. Holotype g, QUEENSLAND: Palmerston (DEI, Eberswalde). —
N.T., Qtp. (Probably = malayana).
rufiventris Macquart, 1847 : 96 (80). Holotype 9, ‘Tasmania’ (BMNH, London) [examined].
— QOtp., Tasm. (?).
siberita Fabricius, 1775 : 798 (Stomoxys). Type(s), DENMARK: Copenhagen (lost). — N.S.W.,
Qup.; widespread EuropE and Asia. (Possibly misidentified from Australia)
sibivita. Incorrect subsequent spelling of sibevita Fabricius.
confusa Malloch, 1930a@: 115 (as var. of sibevita). Holotype 3, QUEENSLAND: Eidsvold
(SPHTM, Sydney) [examined].
surda Curran, 1938: 190. Holotype g, QUEENSLAND: Kuranda (BMNH, London, ex coll.
Wainwright) [examined]. — N.S.W., Qip., W.A.

116 R. W. CROSSKEY

tenuis Malloch, 1930a: 114. Holotype g, NEw SoutH Waters: Gundamain National Park
(SPHTM, Sydney) [examined]. — N.S.W., QLD. (?).

varia Curran, 1929: 509. Holotype g, NEw SoutH WaALEs (not located). —N.S.W.

variegata Curran, 1929: 509. Holotype g, QUEENSLAND: Gravesend (SPHTM, Sydney)
[examined]. — OLD.

vittata Guérin-Méneville, 1838: 299. Holotype 9, NEw SoutH Wates: Port Jackson
(=Sydney) (MNHN, Paris) [examined]. — N.S.W.

vittata Macquart, 1843 : 249 (92). Holotype 2, New Sours WatEs: Port Jackson (=Sydney)

(MNHN, Paris) [examined]. Junior primary homonym and junior objective synonym
of P. vittata Guérin-Méneville.

It is considered certain (Crosskey, 1971 : 284-285) that Macquart described his P.
vittata from the same female specimen from Port Jackson that Guérin-Méneville had already
used as the basis of his P. vittata, Macquart apparently being unaware of Guérin-Méneville’s
description. P. vittata Macquart has therefore the status of a junior objective synonym
and junior homonym of P. vittata Guérin-Méneville.

Macquart (1843 : 249) considered that vittata might be the female of P. conica Guérin-
Méneville. As the male holotype of conica and the female holotype of vittata (both from
Port Jackson) are mounted and labelled exactly similarly it is almost certain that they
were collected at the same time, and probable that the sexes associate. Synonymy is
not established at present.

Genus PROSENINA Malloch

Prosenina Malloch, 1930a : 116. Type-species: Prosenina nicholsoni Malloch, 1930, by original
designation.

nicholsoni Malloch, 1930a: 116. Holotype g, NEw SoutH Wates: Gundamain National
Park (ANIC, Canberra) [examined]. — N.S.W., Tasm., W.A.

Genus RUTILOTRIXA Townsend

Rutilotrixa Townsend, 1933 : 448. Type-species: Tvixva lateralis Walker, 1849, by original
designation.

Ruya Paramonov, 1968 : 381. Type-species: Chaetogastey diversa Paramonov, 1954, by
original designation. Syn. n.

diversa Paramonov, 1954 : 280 (Chaetogastev). Holotype gj, AUSTRALIAN CAPITAL TERRITORY:
Blundell’s (ANIC, Canberra) [examined]. Comb. n.—A.C.T., N.S.W., Vict. (Probably
= lateralis).

lateralis Walker, 1849 : 698 (Trixa). Holotype 9, [AustraLta] (BMNH, London) [examined].
— AUSTRALIA (state unknown, probably N.S.W.).

No locality was given in the original description; Townsend (1933 : 448) rightly cited

Australia from evidence at the BMNH.

monstruosa Paramonov, 1968 : 383 (Ruwya). Holotype 9, NEw SoutH Wates: Buraja
(ANIC, Canberra). Comb. n.—N.S.W.

westralica Paramonov, 1968 : 382 (Rwya). Holotype g, WESTERN AUSTRALIA: Pinjarra
(ANIC, Canberra). Comb. n.— W.A.

Genus SENOSTOMA Macquart

Senostoma Macquart, 1847 : 96 (80). Type-species: Senostoma variegata Macquart, 1847, by
monotypy.

Rhynchiodexia Bigot, 1885a:xi. Type-species: Rhynchiodexia tenuipes Bigot, 1885, by
monotypy. (NEWCALEDONIA). Syn. n.

SS

TACHINIDAE OF AUSTRALIA 117

Austrodexia Malloch, 1930a@:122. Type-species: Austrodexia setigera Malloch, 1930, by
original designation. Syn. n.

Lasiocalypter Malloch, 1930a: 119. Type-species: Lasiocalypter flavohirta Malloch, 1930, by
original designation. Syn. n.

Lasiocalyptrina Malloch, 1930a : 122. Type-species: Lasiocalyptrina modesta Malloch, 1930,
by original designation. Syn. n.

apicale Curran, 1938 ; 193 (Lasiocalypter). Holotype g, NEw Soutu WaLEs: Jenolan caves
(SPHTM, Sydney) [examined]. Comb. n.-—N.S.W.
appendiculatum Macquart, 1851 : 202 (229) (Dexia). Holotype g, ‘Tasmania’ (MNHN,
Paris) [examined]. Comb. n.—N.S.W., Oxp, Taso. (?).
punctum Walker, 1858 : 205. Lectotype ¢ [not 9] (by present designation), NEw SoutH
WALEs (BMNH, London) [examined]. Syn. n.
atripes Malloch, 1930a : 121 (Lasiocalypter). Holotype g, NEw SoutH WateEs: Blue Mts
(SPHTM, Sydney) [examined]. Comb. n.-—N.S.W.
basale Curran, 1938 : 193 (Lasiocalypter). Holotype g, QUEENSLAND: Macpherson Range,
Roberts Plateau (SPHTM, Sydney) [examined]. Comb. n.-QLp.
brevipalpe Macquart, 1846 : 317 (189) (Omalogaster). Lectotype ¢ (by designation of Crosskey,
1971 : 280), TasMANIA (MNHN, Paris) [examined]. Comb. n.— Tas.
brevipalpe Rondani, 1864: 22 (Dexia). Type(s), AUSTRALIA (not located, probably lost).
Comb. n. — AustrRAtia (state unknown).
S. brevipalpe Rondani is a junior secondary homonym of S. brevipalpe Macquart. No
new name is proposed at the present time.
commune Malloch, 1930a@ : 125 (Austrodexia). Holotype J, AUSTRALIAN CAPITAL TERRITORY:
Canberra (ANIC, Canberra) [examined]. Comb. n.—A.C.T., N.S.W., OLD.
flavohirtum Malloch, 1930a: 121 (Lasiocalypter). Holotype g, NEw SoutH WALEs:
Barrington Tops (SPHTM, Sydney) [examined]. Comb. n.—N.S.W.
hirticauda Malloch, 1930a : 120 (Lasiocalypter). Holotype g, NEw Soutu Wa LEs: Barrington
Tops (SPHTM, Sydney) [examined]. Comb. n.—N.S.W.
hyria Walker, 1849 : 843 (Dexia). Type(s), SourH AusTRALta: Adelaide (lost). Comb. n.—
S.A.
mixtum Malloch, 1930a : 126 (Austrodexia). Holotype g, NEw Soutn WaLEs: Barrington
Tops (SPHTM, Sydney) [examined]. Comb. n.—N.S.W.
modestum Malloch, 1930a@ : 122 (Lasiocalyptrina). Holotype g, Victoria: Gisborne (SPHTM,
Sydney) [examined]. Comb. n.—N.S.W., Tasm., Vict.
nigrihirtum Malloch, 1930a : 119 (Lasiocalypter). Holotype g, NEw SoutuH WALEs: Kosciusko
{publ. as ‘Barrington Tops’] (SPHTM, Sydney) [examined]. Comb. n.-—N.S.W., Vict.
notatum Walker, 1852 : 309 (Dexia). Holotype 9, NEw SoutH Wares (BMNH, London)
[examined]. Comb. n.-—N.S.W., Vict.
pallidihirtum Malloch, 1930a : 126 (Austrodexia). Holotype g, NEw Soutu WatEs: Barring-
ton Tops (SPHTM, Sydney) [examined]. Comb. n.—N.S.W.
punctipenne Macquart, 1846 : 315 (187) (Dexia). Holotype gj, AustrALIaA (BMNH, London)
[examined]. Comb. n.— AusTRALIA (state unknown).
pictipennis. Incorrect subsequent spelling (lapsus for punctipennis by Malloch, 1930a : 123).
Malloch (1930a@ : 124) reports this species from N.S.W. and W.A. but his identification
(as ‘pictipennis’) needs confirmation.
rubricarinatum Macquart, 1846 : 315 (187) (Dexia). Holotype g, Tasmania (MNHN,
Paris) [examined]. Comb. n.—Tasm.
setigerum Malloch, 1930a : 124 (Austrodexia). Holotype g, NEw SoutH WaLEs: Woy Woy
(SPHTM, Sydney) [examined]. Comb. n.-—N.S.W., OLp.
Setiventre Malloch, 1930a : 126 (Austrodexia). Holotype g, NEw SoutH WatEs: Glenreagh
(SPHTM, Sydney) [examined]. Comb. n.—N.S.W., OLp.
taylori Curran, 1938 : 191 (Austrodexia). Holotype 9, QuEENSLAND: Kuranda (SPHTM,
Sydney) [examined]. Comb. n.- QLp.

118 R. W. CROSSKEY

tessellatum Macquart, 1851 : 202 (229) (Dexia). Holotype 9, ‘Tasmania’ (MNHN, Paris)
[examined]. Comb. n.-— Taso. (?).

testaceicorne Macquart, 1851 : 201 (228) (Dexia). Holotype g, TASMANIA [publ. as ‘Oceania’ |
(MNHN, Paris) [examined]. Comb. n.—Tasm.

unipunctum Malloch, 1930a : 126 (Austrodexia). Holotype g, NEw SoutH WALEs: Barrington
Tops (SPHTM, Sydney) [examined]. Comb. n.—N.S.W.

variegatum Macquart, 1847 : 96 (80) (as variegata). Holotype 9, Tasmania (BMNH, London)
[examined]. — N.S.W., Tasm.

Genus TRICHOSTYLUM Macquart

Trichostylum Macquart, 1851 : 181 (208). Type-species: Tvichostylum vufipalpis Macquart,
1851, by monotypy.

rufipalpe Macquart, 1851 : 182 (209) (as rufipalpis). Holotype 2, AustrRaL1a (MNHN, Paris)
[examined]. — AUSTRALIA (state unknown; publ. as ‘cote orientale’, prob. N.S.W. or QLD).

Tribe RUTILIINI Brauer & Bergenstamm

RUTILIIDAE Brauer & Bergenstamm, 1889: 76, 152. Type-genus: Rutilia Robineau-
Desvoidy, 1830.

Genus AMPHIBOLIA Macquart

Amphibolia Macquart, 1843 : 278 (121). Type-species: Amphibolia valentina Macquart, 1843,
by original designation.

Subgenus AMPHIBOLIA Macquart

Amphibolia Macquart, 1843 : 278 (121). Type-species: Amphibolia valentina Macquart, 1843,
by original designation.

albocincta Malloch, 1930@ : 108 (Rutilia). Holotype 9, NEw SoutH Wates: Barrington
Tops (ANIC, Canberra) [examined]. — A.C.T., N.S.W.

campbelli Paramonov, 1950 : 523. Holotype g, AUSTRALIAN CAPITAL TERRITORY: Blundell’s
(ANIC, Canberra) [examined]. — A.C.T., N.S.W., Vict.

commoni Paramonov, 1968 ; 363. Holotype 9, Vicror1a: Grampians, Fyan’s Creek (ANIC,
Canberra). — N.S.W., Vict.

ignorata Paramonov, 1950 : 522. Holotypeg, New Soutu Wa tEs: Tindery (ANIC, Canberra)
[examined]. — A.C.T., N.S.W., S.A., Vict., W.A.; Lorp Howe I.

valentina Macquart, 1843 : 279 (122). Type(s) g, AUsTRALIA (MHN, Lille).-N.S.W., QLp.,
Tasm., Vict., W.A.

vidua Guérin-Meéneville, 1843 : 273 (Rutilia). Syntypes (1 ¢, 3 9), AUSTRALIA (lost).

wilsoni Paramonov, 1950: 524. Holotype g, VicroriaA: Warburton (NMV, Melbourne). —

VICT.

Subgenus PARAMPHIBOLIA Brauer & Bergenstamm

Pavamphibolia Brauer & Bergenstamm, 1891 : 389 (85) (as genus). Type-species: Rutilia
assimilis Macquart, 1851, by monotypy.

Chaetogastrina Malloch, 1929b: 313. Type-species: Chaetogastrina stolida Malloch, 1929, by
original designation.

assimilis Macquart, 1851 : 192 (219) (Rutilia). Lectotype ¢ (by designation of Crosskey,
1971 : 285), TASMANIA (MNHN, Paris) [examined]. — Tasm., Vict.

stolida Malloch, 1929b : 313 (Chaetogastrina). Holotype g, NEw SoutH Wa tes: Barrington
Tops (AM, Sydney) [examined]. — N.S.W.

pi

| -—a

TACHINIDAE OF AUSTRALIA 119

Genus CHETOGASTER Macquart

Chetogaster Macquart, 1851 : 198 (225). Type-species: Chetogaster violacea Macquart, 1851,
by monotypy.

Codium Enderlein, 1936: 417. Type-species: Rutilia oblonga Macquart sensu Enderlein
(misidentification) [= Chetogaster violacea Macquart], by original designation.

Chaetogaster. Incorrect subsequent spelling of Chetogastey Macquart.

argentifera Malloch, 1936: 19. Holotype 2, Victoria: Gisborne (SPHTM, Sydney) [examined].
Oe Glu NS. WV... VICE:
auriceps Paramonov, 1968 : 371. Holotype 2, QUEENSLAND: Collinsville (ANIC, Canberra). —
OLD.
canberrae Paramonoy, 1954: 277. Holotype g, AUSTRALIAN CapiTAL TERRITORY: Black
Mt (ANIC, Canberra) [examined]. —A.C.T., N.S.W., Qup, Vict.
oblonga Macquart, 1847: 92 (76) (Rutilia). Holotype g, Austratia (BMNH, London,
{examined]. —- N.S.W., Vict.
gvatiosa Paramonov, 1954 : 283. Holotype g, Vicrorta: Gibraltar Point, Wonnangatta
River (ANIC, Canberra) [examined].
pellucida Paramonov, 1954 : 276 (as var. of argentifera). Holotype g, NEw SoutH WALES:
Toronto (ANIC, Canberra) [examined]. — N.S.W.
violacea Macquart, 1851 : 198 (225). Holotype g, AusTRALIA (MNHN, Paris) [examined]. —
PAG. a, Nes. W.; Orp. Vict.
[oblonga Macquart sensu Enderlein (misidentification) |
viridis Malloch, 1936: 19. Holotype g, NEw SoutH Wares: Comboyne (SPHTM, Sydney)
fexamined].— N.S.W., OLp.

Genus CHRYSOPASTA Brauer & Bergenstamm

Chrysopasta Brauer & Bergenstamm, 1889: 152 (84). Type-species: Chrysopasta versicolor
Brauer & Bergenstamm, 1889 [= Rutilia elegans Macquart, 1846], by original designation
and monotypy.

Roederia Brauer & Bergenstamm, 1893 : 98 (10). Type-species: Chrysopasta versicoloy Brauer
& Bergenstamm, 1889 [=Rutilia elegans Macquart, 1846], by monotypy (see Crosskey,
1973 : 103). [Junior homonym of Roederia Mik, 1881.]

Echrysopasta Townsend, 1932 : 39. Type-species: Rutilia elegans Macquart, 1846, by original
designation.

Euchrysopasta. Incorrect subsequent spelling of Echrysopasta Townsend (Paramonov, 1968).

elegans Macquart, 1846 : 309 (181) (Rutilia). Holotype g, [WESTERN AusTRALIA] (BMNH,
London) [examined]. — W.A.
Type-locality originally cited as Sydney, New South Wales, in error (Crosskey, 1973 : 106).
zabiyna Walker, 1849 : 863 (Dexia). Lectotype ¢ (by designation of Crosskey, 1973 : 118),
WESTERN AUSTRALIA: Perth (BMNH, London) [examined].
versicolor Brauer & Bergenstamm, 1889 : 171 (103). Lectotype ¢ (by fixation of Malloch,
1928a : 616), WESTERN AUSTRALIA: Swan River (NM, Vienna).
zabryina. Incorrect subsequent spelling of zabirna Walker.

Genus FORMOSIA Guérin-Méneville
Formosia Guérin-Méneville, 1843 : 263. Type-species: Rutilia mirabilis Guérin-Méneville,
1831, by monotypy. (NEw GUINEA).
Subgenus EUAMPHIBOLIA Townsend

Euamplibolia Townsend, 1916b: 618 (as genus). Type-species: Rutilia fulvipes Guérin-
Méneville, 1843 [=Rutilia speciosa Erichson, 1842], by original designation.

120 R. W. CROSSKEY

Hega Enderlein, 1936 : 419, 421. Type-species: Hega viridicingens Enderlein, 1936 [=Rutilia
complicita Walker, 1861], by original designation. (Motuccas).

faceta Enderlein, 1936 : 422 (Hega). Holotype g, QUEENSLAND: North Queensland (MNHU,
Berlin) [examined]. — QLp.
smaragdina Malloch, 1929b : 312. Holotype 9, QUEENSLAND: Gordonvale (AM, Sydney)
[examined]. — QLD.
speciosa Erichson, 1842 : 273 (Rutilia). Lectotype 9 (by designation of Crosskey, 1973 : 120),
Tasmania (MNHU, Berlin) [examined]. —A.C.T., N.S.W., Quip, Tasm., Vict., W.A.
fulvipes Guérin-Méneville, 1843 : 273 (Rutilia). Holotype 9, AustTrRatta (lost).

Genus PRODIAPHANIA Townsend

Diaphania Macquart, 1843 : 277 (120). Type-species: Diaphania testacea Macquart, 1843, by
monotypy. [Junior homonym of Diaphania Hiibner, 1818.]

Prodiaphania Townsend, 1927b : 159. [Replacement name for Diaphania Macquart.]

[Senostoma Macquart sensu authors (misidentification) ]

arida Paramonov, 1968 : 397. Holotype g, Vicrorta: Little Desert, 5 mls S. of Kiata (ANIC,
Canberra). — Vict.

biarmata Malloch, 1936 : 14 (Senostoma). Holotype g, SourH AusTRALIa (SPHTM, Sydney)
[examined].—S.A.

brevitarsis Paramonov, 1968 : 395. Holotype g, NEw SoutTH WALEs: near Queanbeyan
(ANIC, Canberra). — N.S.W.

claripennis Malloch, 1929) : 292 (as var. of testacea). Holotype g, WESTERN AUSTRALIA:
King George’s Sound (AM, Sydney) [examined]. — W.A.

commoni Paramonov, 1968 : 389. Holotype g, Vicrorta: Little Desert, 5 mls S. of Kiata
(ANIC, Canberra). —S.A., Vict.

cygnus Malloch, 1936: 15 (Senostoma). Holotype g, WESTERN AUSTRALIA: Swan River
(SPHTM, Sydney) [examined]. — W.A.

deserta Paramonov, 1968 : 398. Holotype g, NEw SoutH WaLes: Wanaaring (ANIC,
Canberra). — N.S.W., QLD.

echinomides Bigot, 1874: 466 (Rutilia). Holotype 9, AustTraLtia (BMNH, London)
[examined]. - AUSTRALIA (state unknown).

echinomyidea. Incorrect subsequent spelling of echinomides Bigot (Brauer, 1899 : 512).

fullerae Paramonov, 1968 : 393. Holotype fg, NEw Soutu WatEs: Barrington Tops (ANIC,
Canberra). — N.S.W.

funebris Paramonov, 1968 : 391. Holotype g, SouTH AUSTRALIA: 40 mls SW of Iron Knob
(ANIC, Canberra).—S.A., W.A.

Jfurcata Malloch, 1936: 14 (Senostoma). Holotype g, AUSTRALIAN CAPITAL TERRITORY:
Canberra (ANIC, Canberra) [examined]. — A.C.T., N.S.W., S.A., Vict.

genitalis Paramonov, 1968 : 400. Holotype g, QUEENSLAND: Herberton (BMNH, London)
[examined]. —A.C.T., N.S.W., QLp, Vict.

paratestacea Paramonov, 1968 : 397. Holotype g, NEw SoutH WaLEs: Wee Jasper (ANIC,

Canberra).

gZeorgei Malloch, 1929b: 292. Holotype g, WESTERN AUSTRALIA: King George’s Sound
(AM, Sydney) [examined]. — N.S.W., S.A., W.A.

minuta Paramonov, 1968 : 399. Holotype g, QUEENSLAND: Gordonvale (ANIC, Canberra). —
QLD.

regina Malloch, 1936: 14 (Senostoma). Holotype g¢, QUEENSLAND: Ejidsvold (SPHTM,
Sydney) [examined]. — OLp.

testacea Macquart, 1843 : 278 (121) (Diaphania). Type(s), AUSTRALIA (MHN, Lille). — A.C.T.,
N.S.W., QLp, Tasm.

victoriae Malloch, 1936 : 13 (Senostoma). Holotype g, Victoria: Gisborne (SPHTM, Sydney)
[examined]. — N.S.W., QLD, Vict.

TACHINIDAE OF AUSTRALIA 121

vittata Macquart, 1855: 126 (106) (Rutilia). Holotype 9, Sourn AvustTRraLia: Adelaide
(BMNH, London) [examined].—S.A., Tas. (?). (Probably = testacea 9).

walkeri Paramonov, 1968 : 400. Holotype 9, WESTERN AUSTRALIA (AMNH, New York). —
W.A.

Genus RUTILIA Robineau-Desvoidy

Rutilia Robineau-Desvoidy, 1830 : 319. Type-species: Tachina vivipara Fabricius, 1805, by
subsequent designation of Crosskey (19674 : 26).

Subgenus AMENIAMIMA Crosskey

Ameniamima Crosskey, 1973 : 51. Type-species: Rutilia argentifera Bigot, 1874, by original
designation.

argentifera Bigot, 1874 : 464. Lectotype g (by designation of Crosskey, 1971 : 300), NEw
SoutH WaLEs: Sydney (BMNH, London) [examined]. — N.S.W., QLp.
frontosa Brauer & Bergenstamm, 1891 : 418 (114). Nomen nudum.
frontosa Malloch, 1929) : 310 (Formosia). Holotype g, NEw SoutH Wa tes: Jindabyne
(AM, Sydney) [examined].
cingulata Malloch, 1930a : 105 (Formosia). Holotype g, NEw SoutH WaLEs: Wentworth
Falls (ANIC, Canberra) [examined]. — N.S.W.
quadripunctata Malloch, 1930a: 104 (Formosia). Holotype g, QUEENSLAND: Eidsvold
(ANIC, Canberra) [examined]. — QLD, Vict.

Subgenus CHRYSORUTILIA Townsend

Chrysorutiia Townsend, 1915): 23 (as genus). Type-species: Rutilia formosa Robineau-
Desvoidy, 1830, by original designation.

Habrota Enderlein, 1936 : 399. Type-species: Rutilia formosa Robineau-Desvoidy, 1830, by
original designation. [Objective synonym of Chrysorutilia.]

Zoramsceus Enderlein, 1936: 416. Type-species: Rutilia erichsonii Engel, 1925 [=Devia
chersipho Walker, 1849], by original designation.

caeruleata Enderlein, 1936 : 402 (Chrysorutilia). Lectotype ¢ (by designation of Crosskey,
1973 : 117), WESTERN AUSTRALIA: Marloo Station (MNHU, Berlin) [examined]. — W.A.
lineata Enderlein, 1936 : 407 (Chrysorutilia). Lectotype ¢ (by designation of Crosskey,
1973 : 118), WESTERN AUSTRALIA: Wurarga, Marloo Station (MNHU, Berlin) [examined].
caesia Enderlein, 1936: 402 (Chrysorutilia). Lectotype g (by designation of Crosskey,
1973 : 117), AUSTRALIA (MNHU, Berlin) [examined]. — N.T., QLp, W.A.
viridescens Enderlein, 1936 : 403 (Chrysorutilia, as var. of caesia). Holotype 9, QUEENSLAND
(MNHU, Berlin) [examined].
rufibarbis Enderlein, 1936 : 405 (Chrysorutilia). Lectotype ¢ (by designation of Crosskey,
1973 : 118), WESTERN AUSTRALIA: Wurarga, Marloo Station (MNHU, Berlin) [examined].
chersipho Walker, 1849 : 864 (Dexia). Neotype ¢ (by designation of et LO73) > £23);
WESTERN AUSTRALIA: Waroona (BMNH, London) [examined]. — W.A.
evichsonii Brauer & Bergenstamm, 1891 : 418 (114). Nomen nudum.
evichsonti Engel, 1925 : 363. Lectotype Q (by fixation of Malloch, 1929b : 297), WESTERN
AUSTRALIA: Swan River (NM, Vienna) [examined].
evichsoni. Incorrect subsequent spelling of evichsonit Brauer & Bergenstamm.
corona Curran, 1930 : 3. Holotype g, NEw Sout WatEs (AMNH, New York) [examined]. —
N.S.W.

122 R. W. CROSSKEY

cryptica Crosskey, 1973 : 65. Holotype g, SourH AUSTRALIA: near Moonta (BMNH, London)
[examined]. — N.S.W., S.A., Vict.
decora Guérin-Méneville, 1843 : 266. Neotype ¢ (by designation of Crosskey, 1973 : 124),
QUEENSLAND: Burpengary (BMNH, London) [examined]. — N.S.W., QLp, Tasm.
formosa Robineau-Desvoidy, 1830 : 320. Neotype ¢ (by designation of Crosskey, 1973 : 124),
NEw SouTtH WALES: near Lake George (ANIC, Canberra) [examined]. —A.C.T., N.S.W.,
Vict., W.A. (?).
uzita Walker, 1849 : 860 (Dexia). Holotype 9, Austratia (BMNH, London) [examined].
pubicolis Thomson, 1869 : 530. Lectotype g (by designation of Crosskey, 1973 : 120),
NEw SoutH WALEs: Sydney (NR, Stockholm) [examined].
subvittata Malloch, 1929b : 295 (as var. of formosa). Holotype 9, WESTERN AUSTRALIA:
King George’s Sound (AM, Sydney) [examined].
There is some doubt about the placement of this name: the type was seen in 1965 but
was not available during a recent revision of Rutiliini (Crosskey, 1973).
goerlingiana Enderlein, 1936 : 404 (Chrysorutilia). Lectotype 9 (by designation of Crosskey,
1973 : 117), WESTERN AUSTRALIA: Wurarga, Marloo Station (MNHU, Berlin) [examined]. —
W.A.
idesa Walker, 1849 : 858 (Dexia). Holotype g [mot 9], Austrat1a (BMNH, London)
[examined]. — AUSTRALIA (state unknown).
imperialis Guérin-Méneville, 1843 : 265. Neotype ¢ (by designation of Crosskey, 1973 : 125),
NEw Soutu WALES: Mount Wilson (ANIC, Canberra) [examined]. — N.S.W., QLD, Vict.
vuficormis Bigot, 1880 : 88. Holotype g, AusTRALIA (BMNH, London) [examined].
semifulva Bigot, 1880 : 89. Lectotype g (by designation of Crosskey, 1971 : 301), AUSTRALIA
(BMNH, London) [examined].
imperialoides Crosskey, 1973 : 67. Holotype ¢, NEw SoutH WALEs: Wee Jasper (BMNH,
London) [examined]. — A.C.T., N.S.W., QLp, Vict.
panthea Walker, 1849 : 862 (Dexia). Holotype 9, WESTERN AUSTRALIA (BMNH, London)
[examined].—S.A., W.A.
rubriceps Macquart, 1847 : 92 (76). Holotype 9, ‘Tasmania’ (BMNH, London) [examined]. —
QLD, Tas. (?); CEYLON, INDIA, VIETNAM, TIMoR (?).
sevena Walker, 1849 : 865 (Dexia). Neotype 2 (by designation of Crosskey, 1973 : 123),
Inp14: Maharashtra, Purandhar, near Poona (BMNH, London) [examined.]
nitens Macquart, 1851 : 189 (216). Holotype 9, Inp1a (MNHN, Paris) [examined].
formosina Curran, 1930: 2. Holotype g, AusTRALIA (AMNH, New York) [examined].
angustigena Enderlein, 1936 : 403 (Chrysorutilia). Lectotype ¢ (by designation of Crosskey,
1973 : 117), QUEENSLAND: Herberton (MNHU, Berlin) [examined].
splendida Donovan, 1805: plate fig. (unnumbered) & description (unpaginated) (Musca).
Neotype ¢ (by designation of Crosskey, 1973 : 123), QUEENSLAND: Brisbane (BMNH,
London) [examined). — N.S.W., QLpD, Vict.
australasia Gray in Cuvier, 1832 : 793 & Plate 114, fig. 1. Type(s), AUSTRALIA (lost).
confluens Enderlein, 1936 : 407 (Chrysorutilia, as var. of splendida). Lectotype g (by
designation of Crosskey, 1973 : 118), QUEENSLAND (MNHU, Berlin) [examined].
evanescens Enderlein, 1936 : 407 (Chrysorutilia, as var. of splendida). Holotype g, AUSTRALIA
(MNHU, Berlin) [examined].
transversa Malloch, 1936: 15. Holotype g, WESTERN AUSTRALIA: Swan River (SPHTM,
Sydney) [examined]. — W.A.

Subgenus DONOVANIUS Enderlein

Donovanius Enderlein, 1936 : 409 (as genus). Type-species: Musca regalis Guérin-Méneville,
1843, by original designation.

Psaronia Enderlein, 1936 : 414. Type-species: Psaronia bisetosa Enderlein, 1936, by original
designation.

TACHINIDAE OF AUSTRALIA 123

Menevillea Enderlein, 1936 : 416. Type-species: Rutilia pellucens Macquart, 1846, by original
designation.
agalmiodes Enderlein, 1936 : 412 (Donovanius). Holotype gf, QUEENSLAND: Cairns (MNHU,
Berlin) [examined]. — OLD.
analoga Macquart, 1851: 191 (218). Holotype Q [not g], ‘Tasmania’ (MNHN, Paris)
[examined].— A.C.T., N.S.W., Orb, Tas. (?), Vict.
dubitata Malloch, 1929b : 303. Holotype 9, NEw SoutH WatgEs: Jindabyne (AM, Sydney)
[examined].
bisetosa Enderlein, 1936: 414 (Psavronia). Lectotype Q (by designation of Crosskey,
1973 : 119), WESTERN AUSTRALIA: Wurarga, Marloo Station (MNHU, Berlin) [examined]. —
W.A.
nigvibasis Enderlein, 1936 : 411 (Donovanius, as var. of fulgidus). Lectotype ¢ (by designation
of Crosskey, 1973 : 118), WESTERN AUSTRALIA: Wurarga, Marloo Station (MNHU,
Berlin) [examined].
ethoda Walker, 1849 : 856 (Dexia). Holotype 9, WESTERN AUSTRALIA: Swan River (BMNH,
London) [examined]. — W.A.
inusta Wiedemann, 1830 : 306 (Tachina). Lectotype 9 (by designation of Crosskey, 1973 : 121)
NEw SoutH WALEs: Port Jackson (=Sydney) (MNHU, Berlin) [examined].—A.C.T.,
N.S.W., QLp, S.A., Tasm., Vict., W.A. (?).
potina Walker, 1849 : 857 (Dexia). Holotype 9, Tasmania (BMNH, London) [examined].
spinipectus Thomson, 1869 : 530. Lectotype ¢ (by designation of Crosskey, 1973 : 120),
NeEw Soutu WaALEs: Sydney (NR, Stockholm) [examined].
castanipes Bigot, 1880 : 87. Lectotype 9 (by designation of Crosskey, 1971 : 300), AUSTRALIA
(BMNH, London) [examined].
castanifrons Bigot, 1880 : 88. Holotype 9, AustraL1a (BMNH, London) [examined].
lepida Guérin-Méneville, 1843 : 268. Neotype ¢ (by designation of Crosskey, 1973 : 125),
AUSTRALIAN CAPITAL TERRITORY: Blundell’s (ANIC, Canberra) [examined].—A.C.T.,
N.S.W., Vict.
fulgida Macquart, 1846 : 308 (180). Lectotype g (by designation of Crosskey, 1971 : 286),
New Soutu WALEs: Sydney (BMNH, London) [examined].
onoba Walker, 1849 : 859 (Dexia). Holotype g, AustRALIa (BMNH, London) [examined].
pellucens Macquart, 1846 : 305 (177). Neotype 3 (by designation of Crosskey, 1973 : 126),
NEw SouTH WALEs: Durras Bay (BMNH, London) [examined]. — A.C.T., N.S.W., Vict.
imitatoy Enderlein, 1936: 412 (Donovanius). Holotype g, AusTRALIA (MNHU, Berlin)
[examined].
regalis Guérin-Méneville, 1831 : Plate 21, fig. 1, rA—-E; 1838 : 295. Neotype g (by designation
of Crosskey, 1973 : 126), AUSTRALIAN CAPITAL TERRITORY: Tharwa (ANIC, Canberra)
[examined]. — A.C.T., N.S.W., S.A., Vict.
nigra Macquart, 1846 : 305 (177). Nomen nudum.
retusa Fabricius, 1775 : 775 (Musca). Holotype 9, AusTRaLIA (BMNH, London: Banks
Coll.) [examined].—W.A. (Type-locality unknown, probably not W.A.).
aditha Walker, 1849: 854 (Dexia). Lectotype ¢ (by present designation), WESTERN
AUSTRALIA: Swan River (BMNH, London) [examined].
viriditestacea Macquart, 1851: 190 (217). Lectotype g (by designation of Crosskey,
1971 : 290), ‘Tasmania’ (MNHN, Paris) [examined].
sabrata Walker, 1849 : 855 (Dexia). Holotype 9, AusTRALIA (BMNH, London) [examined]. —
N.S.W., QLD.
[inornata Guérin-Méneville sensu Malloch (misidentification) ]
spinolae Rondani, 1864 : 23. Type(s), AUSTRALIA (not located, probably lost).
viridinigra Macquart, 1846 : 307 (179). Lectotype 9 (by designation of Crosskey, 1971 : 290),
‘TASMANIA’ (MNHN, Paris) [examined].—N.S.W., QLp, Taso. (?).
fuscotestacea Macquart, 1846 : 306 (178). Holotype 9, NEw Sout Wa Es: Sydney (BMNH,
London) [examined].
barcha Walker, 1849 : 857 (Dexia). Holotype 9, AustraLia (BMNH, London) [examined].

124 R. W. CROSSKEY

Subgenus GRAPHOLOSTYLUM Macquart

Grvapholostylum Macquart, 1851 : 196 (223) (as genus). Type-species: Grapholostylum dorso-
maculatum Macquart, 1851, by monotypy.

Agalmia Enderlein, 1936 : 433. Type-species: Rutilia albopicta Thomson, 1869 [=Grapholo-
stylum dorsomaculatum Macquart, 1851], by original designation. [Junior homonym of
Agalmia Enderlein, 1934 (Muscidae)].

albovirida Malloch, 1929b: 307. Holotype 9, QUEENSLAND: Yeppoon (AM, Sydney)
[examined]. — QLD.
dorsomaculata Macquart, 1851 : 196 (223) (Grapholostylum). Lectotype ¢ (by designation
of Crosskey, 1971 : 271), ‘TAsMaANIA’ (MNHN, Paris) [examined]. —- N.S.W., Taso. (?).
leucosticta Schiner, 1868 : 319. Holotype 9, [AusTRALIA: publ. as New Zealand in error]
(NM, Vienna) [examined].
albopicta Thomson, 1869 : 529. Holotype 29, NEw SoutH WALEs: Sydney (NR, Stockholm)
[examined].
variegata Bigot, 1874 : 461 (Formosia). Lectotype 9 (by designation of Crosskey, 1971 : 299),
AUSTRALIA (BMNH, London) [examined].
fuscisquama Malloch, 19304 : 107 (as var. of leucosticta). Holotype 9, NEw SoutH WatEs:
Barrington Tops (ANIC, Canberra) [examined].
micans Malloch, 1929b : 299. Holotype 9, NEw SoutH WALEs: Kosciusko (AM, Sydney)
[examined]. — A.C.T., N.S.W.
subtustomentosa Macquart, 1851 : 191 (218). Holotype g, Tasmanta (MHNH, Paris)
[examined]. —- TAsM.
velutina Bigot, 1874 : 463 (Formosia). Lectotype 9 (by designation of Crosskey, 1971 : 299),
TasMANIA (BMNH, London) [examined].

Subgenus MICRORUTILIA Townsend

Microrutiia Townsend, 1915) : 23 (as genus). Type-species: Rutilia minor Macquart, 1846,
by original designation.

Prosenostoma Townsend, 1932 : 39. Type-species: Senostoma flavipes Brauer & Bergenstamm
sensu Townsend (misidentification) [=Rutilia (Senostoma) hirticeps Malloch, 1929], by
original designation.

Eucompsa Enderlein, 1936 : 400. Type-species: Rutilia minor Macquart, 1846, by original
designation. [Junior homonym of Eucompsa Enderlein, 1922 (Tabanidae) and junior
objective synonym of Microrutilia Townsend. ]

Pogonagalmia Enderlein, 1936 : 435. Type-species: Rutilia (Senostoma) hirticeps Malloch,
1929, by original designation. [Junior objective synonym of Prosenostoma Townsend. ]}

[Senostoma Macquart sensu authors (misidentification) ]

agalmiodes Enderlein, 1936 : 434 (Prosenostoma, as aberration of ruficorne). Aberrational
name without status in nomenclature.
cupreiventris Malloch, 1936: 18 (as var. of vuficornis). Holotype g, NEw SouTtH WALEs:
Barrington Tops (SPHTM, Sydney) [examined]. — N.S.W.
fulviventris Bigot, 1874 : 465. Lectotype 9 (by designation of Crosskey, 1971 : 301), TASMANIA
(BMNH, London) [examined]. — Tasm.
flavipes Brauer & Bergenstamm, 1889 : 126 (58) (Senostoma). Lectotype 9 (by designation
of Crosskey, 1973 : 121), AUSTRALIA (NM, Vienna) [examined].
hirticeps Malloch, 1929) : 305. Holotype g, NEw SoutH WateEs: Monaro, Moonbar (AM,
Sydney) [examined]. — N.S.W., Vict., W.A.
pallens Curran, 1930: 2. Holotype g, New SoutH WaLEs (AMNH, New York) [examined].
[flavipes Brauer & Bergenstamm sensu Townsend (misidentification) ]
liris Walker, 1849 : 882 (Musca). Holotype 9 [AusTRALIA, prob. TasMANIA] (BMNH,London)
[examined].—Tasm. (Probably = minor 9).
livis. Incorrect subsequent spelling of livis Walker (Malloch, 19280 : 660).

TACHINIDAE OF AUSTRALIA 125

media Macquart, 1846 : 310 (182). Lectotype ¢ (by designation of Crosskey, 1971 : 286),

TasMANIA (MNHN, Paris) [examined]. —N.S.W., Tasm., Vict.
vuficovnis Macquart, 1851 : 193 (220) (Diaphania). Holotype g, TAsmMAniA (MNHN, Paris)

[examined].

minor Macquart, 1846 : 310 (182). Lectotype ¢ (by designation of Crosskey, 1971 : 286),
TasMANIA (MNHN, Paris) [examined].—Tasm., N.S.W.

nigriceps Malloch, 1929) : 306. Holotype g, NEw SoutH Wa tes, East Dorrigo, Ulong
(AM, Sydney) [examined]. — N.S.W.

nigripes Enderlein, 1936 : 435 (Prosenostoma). Lectotype ¢ (by designation of Crosskey,
1973 : I19), QUEENSLAND: Herberton (MNHU, Berlin) [examined]. — A.C.T., QLp.

Subgenus NEORUTILIA Malloch

Neorutiia Malloch, 1936: 17. Type-species: Rutilia (Neorutilia) simplex Malloch, 1936, by
original designation.

simplex Malloch, 1936:17. Holotype g, QUEENSLAND: Ejidsvold (SPHTM, Sydney)
[examined]. — N.S.W., QLp, Vict.

Subgenus RUTILIA Robineau-Desvoidy

Rutilia Robineau-Desvoidy, 1830 : 319 (as genus). Type-species: Tachina vivipara Fabricius
1805, by subsequent designation of Crosskey (19674 : 26).

Psavoniella Enderlein, 1936: 417. Type-species: Rutilia castanipes Bigot sensu Enderlein
(misidentification) [= Rutila setosa Macquart, 1847], by original designation.

Stivaulax Enderlein, 1936 : 428. Type-species: Tachina vivipara Fabricius, 1805, by original
designation. [Junior objective synonym of Rutilia.]

confusa Malloch, 1929) : 309 (Formosia). Holotype g, SoutH AusTRALIA: Kangaroo Island,
deep creek 20 mls from Kingscote (AM, Sydney) [examined]. — A.C.T., N.S.W., S.A., Vict.
dentata Crosskey, 1973 : 81. Holotype g, Vicrorta: Monbulk (BMNH, London) [examined]. —
Vict.
setosa Macquart, 1847 : 94 (78). Neotype ¢ (by designation of Crosskey, 1973 : 126), NEw
SoutH WALEs: 4 mls N. of Bateman’s Bay (ANIC, Canberra) [examined]. — N.S.W., Vict.
[castanipes Bigot sensu Enderlein (misidentification) ]
vivipara Fabricius, 1805 : 309 (Tachina). Neotype ¢ (by designation of Crosskey, 1973 : 126),
NEw SoutH WALEs: Barrington Tops (ANIC, Canberra) [examined].—A.C.T., N.S.W.,
Qxp, TasM., VIcT.
durvillei Robineau-Desvoidy, 1830 : 321. Neotype (by designation of Crosskey, 1973 : 124),
New Soutu WatLeEs: Hawkesbury River (BMNH, London) [examined].
inornata Guérin-Méneville, 1843 : 268. Lectotype 2 (by designation of Crosskey, 1973 : 119),
AUSTRALIA (MNHN, Paris) [examined].
desvoidyi Guérin-Méneville, 1843 : 269. Lectotype 9 (by designation of Crosskey, 1973 : 119),
AUSTRALIA (MNHN, Paris) [examined].

Subgenerically unplaced species of Rutilia s.1.

micropalpis Malloch, 1929b : 298. Holotype 9, NEw SoutH WaALEs: Como (AM, Sydney). —
N.S.W.

scutellata Enderlein, 1936: 405 (Chrysorutiia, as var. of media). Holotype 9, SouTH
AustRALIA: Adelaide (MNHU, Berlin).—S.A. (Status uncertain)

Unplaced names of Rutiliini

accedens Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum.
accedens Brauer, 1899 : 512 (Rutilia). Unavailable.

126 ROW. CROSSKEY

evvonea Paramonov, 1968 : 356, 361 (Rutilia). Nomen nudum.

grisea Brauer & Bergenstamm, 1891 : 417 (113) (Diaphania). Nomen nudum.
humeralis Paramonov, 1968 : 355 (Rutilia). Nomen nudum.

incomparabilis Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum.
sovoy Brauer & Bergenstamm, 1891 : 418 (114) (Rutilia). Nomen nudum.
viyidithorax Bigot, 1874 : 457 (Formosia). Nomen nudum.

Subfamily TACHININAE Robineau-Desvoidy
TACHINARIAE Robineau-Desvoidy, 1830 : 185. Type-genus: Tachina Meigen, 1803.

Tribe PALPOSTOMATINI Townsend

PALPOSTOMATINI Townsend, 1925: 250. Type-genus: Palpostoma Robineau-Desvoidy,
1830.

Genus APALPOSTOMA Malloch

Apalpostoma Malloch, 1930a: 134. Type-species: Apalpostoma cinerea Malloch, 1930, by
original designation.

cinereum Malloch, 1930a : 134 (as cinerea). Holotype 9, WESTERN AUSTRALIA: Wyalkatchem
(SPHTM, Sydney) [examined]. — W.A.

Genus EUSTACOMYIA Malloch

Eustacomyia Malloch, 1927 : 337. Type-species: Eustacomyia breviseta Malloch, 1927, by
original designation.

breviseta Malloch, 1927 : 337. Holotype g, NEw Soutu WatEs: Sydney (SPHTM, Sydney)
[examined]. — N.S.W.

hirta Malloch, 1930a : 133. Holotype g, NEw Soutu Wates: Killara, Allowrie (SPHTM,
Sydney) [examined]. — N.S.W.

Genus PALPOSTOMA Robineau-Desvoidy

Palpostoma Robineau-Desvoidy, 1830: 429. Type-species: Palpostoma testacea Robineau-
Desvoidy, 1830, by monotypy.

Opsophasiops Townsend, 1915b: 22. Type-species: Myiophasia flava Coquillett, 1900, by
original designation.

Pseudopalpostoma Townsend, 1926c : 533. Type-species: Palpostoma desvoidyi Aldrich, 1922,
by original designation.

aldrichi Hardy, 1938:57. Lectotype g (by present designation), QUEENSLAND: Cairns

(USNM, Washington) [examined]. — QLD.
[testacea Robineau-Desvoidy sensu Aldrich (misidentification) ]

apicale Malloch, 1927 : 339 (as apicalis). Holotype 9, NEw SoutH Wares: Woy Woy
(SPHTM, Sydney) [examined]. — N.S.W.

armiceps Malloch, 1931 : 296. Holotype 2, QUEENSLAND: Townsville (USNM, Washington)
[examined]. — OLD.

desvoidyi Aldrich, 1922: 5. Holotype g, QUEENSLAND: Cairns (USNM, Washington)
[examined]. — A.C.T., QLD.

TACHINIDAE OF AUSTRALIA 127

flavum Coquillett, 1900 : 390 (Myiophasia). Lectotype ¢ (by fixation of Aldrich, 1922: 5

as ‘Type’), WESTERN AUSTRALIA (USNM, Washington) [examined]. — W.A.
Lectotype is labelled as from “Tasmania’ but species was described from ‘West Australia’.

The host beetle from which the type-material was reared is common in Western Australia
but absent from Tasmania.

subsessile Malloch, 1931 : 297 (as subsessilis). Holotype 9 [badly damaged], New Soutu
Wates: Sydney (SPHTM, Sydney) [examined]. — A.C.T., N.S.W. (probably = testacewm)

testaceum MRobineau-Desvoidy, 1830: 429 (as testacea). Type(s), AUSTRALIA (lost). —
AUSTRALIA (probably N.S.W.).

Tribe MYIOTRIXINI Townsend

MYIOTRIXINI Townsend, 1936: 17, 90. Type-genus: Myiotrixa Brauer & Bergenstamm,
1893.

Genus MYIOTRIXA Brauer & Bergenstamm

Myiotrixa Brauer & Bergenstamm, 1893 : 96 (8). Type-species: Myiotrixa prosopina Brauer
& Bergenstamm, 1893, by original designation and monotypy.

prosopina Brauer & Bergenstamm, 1893 : 96 (8). Holotype g, AusTRALIA (NM, Vienna)
[examined]. —- AUSTRALIA (state uncertain).
The provenance is shown on the type label as ‘Austra. sept.’ (i.e. northern Australia)
but was published in the original description as “‘W. Australien’. No other specimens
are known and the provenance remains uncertain.

Tribe ORMIINI Townsend
ORMIINAE Townsend, t915a@ : 53. Type-genus: Ormia Robineau-Desvoidy, 1830.

Genus THEROBIA Brauer

Therobia Brauer, 1862 : 1231. Type-species: Tyypoderma abdominalis Wiedemann, 1830, by
monotypy. (‘BENGAL’).

Therobiopsis Townsend, 1919b : 166. Type-species: Aulacephala braueri Kertesz, 1899, by
original designation. (NEW GUINEA).

Ormiominda Paramonov, 1955 : 125. Type-species: Ormiominda rieki Paramonov, 1955, by
original designation.

rieki Paramonov, 1955 : 126 (Ovmiominda). Holotype 9, QUEENSLAND: Ayr (ANIC, Canberra)
[examined]. — OLp.

secunda Paramonov, 1955 : 127 (Ormiominda). Holotype g, QUEENSLAND: 10 mls W. of
Collinsville (ANIC, Canberra) [examined]. —- QLD.

Tribe GLAUROCARINI Townsend
GLAUROCARINI Townsend, 1926¢ : 529. Type-genus: Glaurocara Thomson, 1869.

Genus DODDIANA Curran

Doddiana Curran, 1927b : 352. Type-species: Doddiana pallens Curran, 1927, by original
designation.

Semisuturia Malloch, 1927 : 339. Type-species: Semisuturia australis Malloch, 1927, by
original designation.

128 R. W. CROSSKEY

australis Malloch, 1927 : 340 (Semisuturia). Holotype 9, QUEENSLAND: Eidsvold (SPHTM,

Sydney) [examined]. — N.S.W., OLD.
maculiventris Malloch, 19330 : 136 (as var. of australis). Holotype 9, NEw SoutH WALEs:

National Park (SPHTM, Sydney) [examined].

flavifrons Malloch, 19306 : 342. Holotype 9, QuEENSLAND: Eidsvold (SPHTM, Sydney)
[examined]. — OLD.

inermis Malloch, 19330 : 138. Holotype 3g, QUEENSLAND: Innisfail (SPHTM, Sydney)
[examined]. — OLD.

pallens Curran, 1927) : 353. Holotype g, QUEENSLAND: Herberton (DEI, Eberswalde)
[examined]. — OLD.

parviseta Malloch, 19306 : 341. Holotype 9, NEw SoutH Wa ss: Sydney (SPHTM, Sydney)
[examined]. — N.S.W.

Tribe CAMPYLOCHETINI Townsend

CAMPYLOCHETINI Townsend, 1936: 21, 23, 229. Type-genus: Campylocheta Rondani,
1859.

Genus ELPE Robineau-Desvoidy
Elpe Robineau-Desvoidy, 1863 : 488. Type-species: Tachina inepta Meigen, 1824, by original
designation. (GERMANY).
Undetermined spp. — N.S.W., S.A., Tas.

Tribe VORIINI Townsend
VORIINI Townsend, 1912 : 50. Type-genus: Vovia Robineau-Desvoidy, 1830.

Genus HYSTRICOVORIA Townsend

Hystricovoria Townsend, 1928 : 395. Type-species: Hystricovoria bakevi Townsend, 1928, by
original designation. (PHILIPPINES).

Undetermined sp. — W.A.

Genus HYLEORUS Aldrich

Hyleovus Aldrich, 1926: 16. Type-species: Hyleorus furcatus Aldrich, 1926, by monotypy.

furcatus Aldrich, 1926: 16. Holotype 9, QUEENSLAND: Cairns (USNM, Washington)
[examined]. — QLtp; NEw GUINEA.

Genus VORIA Robineau-Desvoidy

Voria Robineau-Desvoidy, 1830 : 195. Type-species: Voria latifrons Robineau-Desvoidy, 1830
[=Tachina ruralis Fallén, 1810], by monotypy. (EUROPE).

ruralis Fallén, 1810 : 265 (Tachina). Lectotype ¢ (by present designation), SWEDEN: Skane,
Esperéd (NR, Stockholm) [examined].—N.S.W.; NEw GuINEA; widespread EURASIA,
NortH AMERICA, MEXICO, SOUTH AMERICA.

Identification of Australian specimens as ruvalis correct on present evidence but
confirmation needed.

TACHINIDAE OF AUSTRALIA 129

Tribe THELAIRINI Lioy
THELAREINI Lioy, 1864 : 65. Type-genus: Thelaiva Robineau-Desvoidy, 1830.

Genus HALYDAIA Egger

Halydaia Egger, 1856: 383. Type-species: Halydaia aurea Egger, 1856, by subsequent
designation of Brauer (1893 : 498). (AusTRIA).

Anaperistommyia Townsend, 1926a:15. Type-species: Anaperistommyia optica Townsend,
1926, by original designation. (SUMATRA).

Macropia Malloch, 1930b : 322. Type-species: Macropia rufiventris Malloch, 1930, by original
designation.

Halidaya. Incorrect subsequent spelling of Halydaia Egger, 1856. [Not Halidaya Rondani,
1856 (Sepsidae) ]

mackerrasi Paramonov, 1960 : 699 (Halidaya). Holotype J, QUEENSLAND: Palm Is. (ANIC,
Canberra) [examined]. — QLD.

norrisi Paramonov, 1960 : 698 (Halidaya). Holotype g, AUSTRALIAN CAPITAL TERRITORY:
Canberra (ANIC, Canberra) [examined]. — A.C.T., QLD.

rufiventris Malloch, 1930b : 322 (Macyopia). Holotype g, NEw SoutH Wates: Sydney
(SPHTM, Sydney) [examined]. — N.S.W.

Genus THELAIRA Robineau-Desvoidy

Thelaiva Robineau-Desvoidy, 1830: 214. Type-species: Thelaiva abdominalis WRobineau-
Desvoidy, 1830 [= Musca nigripes Fabricius, 1794], by subsequent designation of Townsend
(1916a : 9). (EUROPE).

Thelairia. Incorrect subsequent spelling of Thelaiva Robineau-Desvoidy (Malloch,
1930@ : I0g-II0).

Theraivia. Incorrect subsequent spelling of Thelaiva R.-D. (Hardy, 1934 : 32-33).

australis Walker, 1852 : 314 (Dexia). Holotype g, AusSTRALIA (BMNH, London) [examined]. —
AUSTRALIA (state unknown).

Undetermined sp. — N.S.W.

[leucozona Panzer sensu Malloch (misidentification)].

Tribe MINTHOINI Brauer & Bergenstamm

MINTHOIDAE Brauer & Bergenstamm, 1889 : 78. Type-genus: Mintho Robineau-Desvoidy,
1830.

Genus MINTHOXIA Mesnil

Minthoxia Mesnil, 1968 : 184. Type-species: Minthoxia dasyops Mesnil, 1968, by original
designation.

dasyops Mesnil, 1968 : 186. Holotype g, New Soutu Wa tes: Lisarow (BMNH, London)
[examined]. — N.S.W.

Genus SUMPIGASTER Macquart

Sumpigastey Macquart, 1855 : 124 (104). Type-species: Sumpigaster fasciatus Macquart, 1855,
by monotypy.

Atractodexia Bigot, 1885b : xxxii. Type-species: Atvactodexia argentifeva Bigot, 1885 [=Sumpi-
gastey fasciatus Macquart, 1855), by monotypy. (NEW CALEDONIA).

I

130 R. W. CROSSKEY

Mesembriomintho Townsend, 1916¢ : 158. Type-species: Mesembriomintho compressa Townsend,
1916 [=Sumpigaster fasciatus Macquart, 1855], by original designation.
fasciatus Macquart, 1855 : 125 (105). Holotype g, QUEENSLAND: Moreton Bay (BMNH,
London) [examined]. — N.S.W., Qrp; NEw CaLeponia, Loyatty Is.
argentifera Bigot, 1885) : xxxii (Atractodexia). Holotype g, NEw CaLreponia (BMNH,
London) [examined].
compressa Townsend, 1916c¢ : 159 (Mesembriomintho). Holotype g. QUEENSLAND: Hamilton,
Upper North Pine (USNM, Washington) [examined].

Tribe NEMORAEINI Robineau-Desvoidy

NEMOREIDAE Robineau-Desvoidy, 1863 (1) : 171. Type-genus : Nemovaea Robineau-
Desvoidy, 1830.

Genus NEMORAEA Robineau-Desvoidy

Nemovaea Robineau-Desvoidy, 1830: 71. Type-species : Nemovaea bombylans Robineau-
Desvoidy, 1830 [= Tachina pellucida Meigen, 1824], by subsequent designation of
Townsend (1916a : 8). (EUROPE).

Undescribed sp. — N.S.W., OLD.

Tribe LESKIINI Townsend
LESKIINI Townsend, 1919c : 20. Type-genus: Leskia Robineau-Desvoidy, 1830.

Genus APATEMYIA Macquart

Apatemyia Macquart, 1846 : 325 (197). Type-species: Apatemyia longipes Macquart, 1846, by
monotypy.

flavipes Macquart, 1851 : 160 (187) (Evorista). Holotype 9, [Tasmanta, publ. as ‘Oceania’]
(MNHN, Paris) [examined]. Comb. n.—Tasm.

longipes Macquart, 1846 : 325 (197). Lectotype ¢ (by designation of Crosskey, 1971 : 263),
TasMANIA (MNHN, Paris) [examined]. — Tasm.

rufiventris Macquart, 1847 : 98 (82) (Calliphorva). Holotype 2, Tasmania (BMNH, London)
[examined]. Comb.n.—Tasm. (Possibly = longipes 9).

Genus DEMOTICOIDES Mesnil

Demoticoides Mesnil, 1953: 150. Type-species: Demoticoides pallidus Mesnil, 1953, by
monotypy. (INDIA).

pallidus Mesnil, 1953 :150. Holotype g, Inp1A: Madras, Nilambur (BMNH, London)
[examined]. — OLp; INp1za.

Genus EXECHOPALPUS Macquart

Exechopalpus Macquart, 1847: 91 (75). Type-species: Exechopalpus rufipalpus Macquart,
1847, by monotypy.

dubitalis Malloch, 1930a: 132. Holotype 3, WESTERN AUSTRALIA: Tammin (SPHTM,
Sydney) [examined]. — W.A.

TACHINIDAE OF AUSTRALIA 131

fulvipes Malloch, 1930a : 132. Holotype 9, WESTERN AUSTRALIA: Eradu (SPHTM, Sydney)
[examined]. — W.A.

nigripes Malloch, 1930a : 132. Holotype 9, New SoutH WaLEs: Sydney (SPHTM, Sydney)
[examined]. — N.S.W.

atripes Malloch, 1930a : 131. [Lapsus for nigvipes in key to species. ]

rufifemur Malloch, 1930a : 131. Holotype g, WESTERN AUSTRALIA: Eradu (SPHTM, Sydney)
[examined]. — W.A.

vufofemorata Malloch, 1930a : 128. [Lapsus for vufifemur in legend, fig. 28.]

rufipalpus Macquart, 1847 : 92 (76). Holotype g, AusTRAL1A (BMNH, London) [examined]. —
AUSTRALIA (state unknown).

vujipalpis. Incorrect subsequent spelling of rufipalpus Macquart (Malloch, 19304 : 130, 132).

Genus RHINOMYOBIA Brauer & Bergenstamm

Rhinomyobia Brauer & Bergenstamm, 1893 : 140 (52). Type-species: Rhinomyobia australis
Brauer & Bergenstamm, 1893, by original designation and monotypy.
Rhinomyiobia. Incorrect subsequent spelling of Rhinomyobia Brauer & Bergenstamm.

australis Brauer & Bergenstamm, 1893 : 140 (52). Holotype 9, AusTRALIA (not located). —
ACL N.S. Ww.

The holotype should be in NM, Vienna, but has not been found in that collection (although
at some time seen there by Townsend).

Genus SIPHOLESKIA Townsend
Sipholeskia Townsend, 1916b : 628. Type-species: Drepanoglossa occidentalis Coquillett, 1895,
by original designation. (NoRTH AMERICA).

certima Curran, 1927) : 351 (Demoticus). WHolotype 9, QUEENSLAND: Kuranda (DEI,
Eberswalde) [examined]. Comb. n.—QLp.

Genus TOXOCNEMIS Macquart

Toxocnemis Macquart, 1855 : 123 (103). Type-species: Tovocnemis vittata Macquart, 1855, by
monotypy.

Toxonemis. Incorrect subsequent spelling of Toxvocnemis Macquart (Hardy, 1934 : 33).

vittata Macquart, 1855: 124 (104). Holotype g, SoutH AustraLia: Adelaide (BMNH,
London) [examined]. — N.S.W., S.A.

Unplaced species of Leskiini

transversalis Malloch, 1930a : 130 (Rhinomyiobia). Holotype 9, QUEENSLAND: Cairns
district (SPHTM, Sydney) [examined]. — QLp.

This species, and allied forms in Fijiand New Guinea, appear to represent an undescribed
genus.

Tribe ERNESTIINI Townsend
ERNESTIINI Townsend, 1912: 50. Type-genus: Evnestia Robineau-Desvoidy, 1830.

132 R. W. CROSSKEY

Genus AMPHITROPESA Townsend

Amphitropesa Townsend, 1933 : 463. Type-species: Amphitropesa elegans Townsend, 1933, by
original designation.

elegans Townsend, 1933 : 464. Holotype 2, NEw SoutH WaLEs (BMNH, London) [examined].
—N.S.W.

Genus CHLOROTACHINA Townsend

Chlovotachina Townsend, 1915b: 21. Type-species: Chrysosoma fiaviceps Macquart, 1851, by
original designation.

Chlorodexia Townsend, 1916¢ : 154. Type-species: Chlovodexia froggattii Townsend, 1916, by
original designation.

flaviceps Macquart, 1851 : 158 (185) (Chrysosoma). Holotype 3, AUSTRALIA (MNHN, Paris)
[examined]. — N.S.W., OLpD, Vict., W.A.

froggattii Townsend, 1916c : 154 (Chlorodexia). Holotype g, NEw SoutH WatEs: Merriwa
(USNM, Washington) [examined]. — N.S.W., OLD.

froggatti. Incorrect subsequent spelling of fvoggattii Townsend (Malloch, 19296 : 324, 326).

nigrocaerulea Malloch, 1929b : 324. Holotype ¢, WESTERN AUSTRALIA: King George’s
Sound (AM, Sydney) [examined]. — W.A., Tas. (?).

Undescribed sp. —S.A., TAsm.

Genus MACROCHLORIA Malloch

Macrochloria Malloch, 19296 : 326. Type-species: Macrochloria calliphorosoma Malloch, 1929
[=Nemoraea nitidiventris Macquart, 1851], by original designation.

nitidiventris Macquart, 1851: 155 (182) (Nemoraea). Holotype g, AusTRALIA (MNHN,
Paris) [examined]. Comb. n.—N.S.W., Tasm.
calliphorosoma Malloch, 19296 : 326. Holotype g, NEw SoutuH WaLEs: Barrington Tops
(AM, Sydney) [examined]. Syn. n.
vufipes Malloch, 1936 : 20 (as var. of calliphorosoma). Holotype g, NEw SoutH WALEs:
Toronto (SPHTM, Sydney) [examined]. Syn. n.

Genus NEXIMYIA Crosskey

Neophasia Brauer & Bergenstamm, 1893 : 100 (12). Type-species: Neophasia picta Brauer
& Bergenstamm, 1893, by original designation and monotypy. [Junior homonym of
Neophasia Behr, 1869. ]}

Euphasia Townsend, 1908: 76. [Replacement name proposed for Neophasia Brauer &
Bergenstamm, but itself a junior homonym of Euphasia Stephens, 1830, and Euphasia
Mulsant & Verreaux, 1876. ]

Neximyia Crosskey, 1967a : 20. [Replacement name for Euphasia Townsend. }

picta Brauer & Bergenstamm, 1893 : 210 (122) (Neophasia). Holotype 9, WESTERN AUSTRALIA
(NM, Vienna) [examined]. — N.S.W., W.A.

Tribe PARERIGONINI Mesnil
PARERIGONINA Mesnil, 1966 : 888. Type-genus: Parerigone Brauer, 1898.

TACHINIDAE OF AUSTRALIA 133

Genus AUSTRALOTACHINA Curran
Australotachina Curran, 1938 : 194. Type-species: Australotachina calliphoroides Curran, 1938,

by original designation.

calliphoroides Curran, 1938 : 195. Holotype g, QUEENSLAND: Kuranda (DEI, Eberswalde)
[examined]. — OLD.

Genus LEVERELLA Baranov
Leverella Baranov, 1934a : 473. Type-species: Leverella institutiimperialis Baranov, 1934, by
original designation. (SOLOMON ISLANDs).
Undetermined sp. — QLD.

Genus PYGIDIMYIA Crosskey
Pygidia Malloch, 1930b : 330. Type-species: Pygidia rufolatevalis Malloch, 1930, by original
designation. [Junior homonym of Pygidia Mulsant & Rey, 1861.]
Pygidimyia Crosskey, 1967a: 25. [Replacement name for Pygidia Malloch.]

rufolateralis Malloch, 1930b : 331 (Pygidia). Holotype g, NEw SoutH Wates: Kosciusko
(ANIC, Canberra) [examined]. — N.S.W.

Genus ZITA Curran

Zita Curran, 1927) : 351. Type-species: Zita aureopyga Curran, 1927, by original designation.

aureopyga Curran, 1927) : 351. Holotype g, QUEENSLAND: Herberton (DEI, Eberswalde)
[examined]. — OLD.

Undetermined sp. — N.S.W.

Tribe LINNAEMYINI Townsend
LINNAEMYINI Townsend, t919a : 591. Type-genus: Linnaemya Robineau-Desvoidy, 1830.

Genus APALPUS Malloch

Apalpus Malloch, 1929) : 318. Type-species: Apalpus dorsalis Malloch, 1929, by original
designation.

dorsalis Malloch, 1929) : 318. Holotype 9, WESTERN AUSTRALIA: Eradu, near Geraldton
(ANIC, Canberra) [examined].—S.A., W.A.

Genus CHAETOPHTHALMUS Brauer & Bergenstamm

Chaetophthalmus Brauer & Bergenstamm, 1891 : 383 (79). Type-species: Micropalpus brevi-
gastey Macquart, 1846, by subsequent designation of Townsend (1916a : 6). [Type-species
not fixed by Brauer & Bergenstamm, 1893; see Crosskey, 19674 : 9.]

Ballardia Curran, 19274 : 166. Type-species: Ballardia pallipes Curran, 1927, by original
designation.

[Amphibolosia Surcouf sensu authors (misidentification) ]

[A protheca Macquart sensu Hardy, 1959 : 215 (misidentification) ]

134 R. W. CROSSKEY

bicolor Macquart, 1848 : 204 (44) (Micropalpus). Holotype 9, AusTRALIA (lost). — N.S.W.,
OLD.

biseriatus Malloch, 1930): 311. Holotype g, NEw SoutH WaLes: Narromine (USNM.
Washington) [examined]. — N.S.W., Orb, S.A.

brevigaster Macquart, 1846: 277 (149) (Micropalpus). Holotype g, Tasmania (BMNH,
London) [examined]. — Tasm., N.S.W. & QLD (?).

pallipes Curran, 1927a : 166 (Ballardia). Holotype ¢ [head lost], QUEENSLAND: Brisbane
(BMNH, London) [examined].—Qtp. (Possibly = bicolor).

ruficeps Macquart, 1847 : 89 (73) (Myobia). Holotype g, Tasmania (BMNH, London)
[examined]. Comb. n.— Tas.

similis Walker, 1852 : 266 (Tachina). Holotype gj, NEw SoutH Wares (BMNH, London)
[examined]. — N.S.W., Vict.

Genus LINNAEMYA Robineau-Desvoidy

Linnaemya Robineau-Desvoidy, 1830:52. Type-species: Linnaemya silvestyis Robineau-
Desvoidy, 1830 [=Tachina vulpina Fallén, 1810], by subsequent designation of Robineau-
Desvoidy (1863 : 131, as vulpina with sylvestris cited in synonymy). (EUROPE).

Linnaemyia. Incorrect subsequent spelling of Linnaemya Robineau-Desvoidy.

concavicornis Macquart, 1851 : 146 (173) (Micropalpus). Holotype 3 [not 9], AusTRALIA
{[prob. NEw SoutH WALES, publ. ‘céte orientale’) (MNHN, Paris) [examined]. Comb. n. —
Orn, NSW. \(?))
nigrvipalpus Tryon, 1900 : 144 (Linnaemyia). Type(s), QUEENSLAND (Not located, possibly
lost). Syn. n.

Tribe TACHININI Robineau-Desvoidy
TACHINARIAE Robineau-Desvoidy, 1830 : 185. Type-genus: Tachina Meigen, 1803.

Genus CUPHOCERA Macquart

Cuphocera Macquart, 1845 : 267. Type-species: Micropalpus ruficornis Macquart, 1835, by
original designation. (EUROPE).

Acuphocera Townsend, 1926a : 37. Type-species: Acuphoceva sumatrensis Townsend, 1926
[=Musca varia Fabricius, 1794], by original designation. (SUMATRA).

emmesia Malloch, 1930b : 318. Holotype g, WESTERN AUSTRALIA: Geraldton (SPHTM,
Sydney) [examined]. — W.A.
pilifacies Macquart, 1851 : 146 (173) (Micropalpus). Holotype ¢ [not 9], AusTRa.ia [prob.
NEw SoutH WALES, publ. ‘céte orientale’) (MNHN, Paris) [examined]. Comb. n.—
N.S.W. (?).
pilosa Malloch, 1930b : 316. Holotype g, NEw SoutH WaLEes: Woy Woy (not located,
probably lost; paratype g in USNM, Washington). —N.S.W.
setigera Malloch, 19300 : 318. Holotype g, NEw Soutu WatEs: Kosciusko (SPHTM, Sydney)
[examined]. — A.C.T., N.S.W.
vittata Macquart, 1846 : 278 (150) (Micropalpus). Holotype g, Tasmania (MNHN, Paris)
[examined]. — Tasm.
varia Fabricius, 1794 : 327 (Musca). Holotype 9, East InpiEes (publ. ‘Ind. or.’) (UZM,
Copenhagen) [examined].— Qtp; NEw GuINEA; widespread ORIENTAL REGION.
sumatrensis Townsend, 1926a : 37. Lectotype ¢ (by designation of Crosskey, 1969 : 90),
SumaTRA: Fort de Kock (ZM, Amsterdam) [examined].
(For other synonyms, based on Oriental types, see Crosskey, 1966a : 674)

TACHINIDAE OF AUSTRALIA 135

Genus ERISTALIOMYIA Townsend

Evistalhomyia Townsend, 1926a : 37. Type-species: Evistaliomyia nitidifrons Townsend, 1926
[=Echinomyia brevipennis Walker, 1857], by original designation. (SUMATRA).
[? Peleteria Robineau-Desvoidy sensu Hardy, 1938 : 66 (misidentification) |

Undescribed sp. — OLD.

Genus MICROTROPESA Macquart

Microtropesa Macquart, 1846 : 313 (185). Type-species: Musca sinuata Donovan, 1805, by
monotypy.

Gerotachina Townsend, 1916c : 152. Type-species: Tachina obtusa Walker, 1852, by original
designation.

Microtropeza. Incorrect subsequent spelling of Microtropesa Macquart.

[Tasmaniomyia Townsend sensu Hardy, 1939 : 33 (misidentification) ]

campbelli Paramonoy, 1951 : 768. Holotype 9, AUSTRALIAN CAPITAL TERRITORY: Blundell’s
(ANIC, Canberra) [examined]. — A.C.T.

canberrae Paramonov, 1951 : 771. Holotype 9, AUSTRALIAN CAPITAL TERRITORY: Canberra,
Black Mt. (ANIC, Canberra) [examined]. — A.C.T.

flavitarsis Malloch, 1929b : 288. Holotype g, Tasmania (AM, Sydney) [examined]. — W.A.,
SA. (?).

flaviventris Malloch, 1930a: 101. Holotype 9, New SoutH Waters: Narromine (ANIC,
Canberra) [examined]. — A.C.T., N.S.W., OLp.

intermedia Malloch, 1930a: 100. Holotype 3, QUEENSLAND: Eidsvold (ANIC, Canberra)
[examined]. — A.C.T., N.S.W., OLp.

latigena Paramonov, 1951 : 769. Holotype 9, WESTERN AusTRALIA: Kalgoorlie (WADA,
Perth).—S.A., W.A.

nigricornis Macquart, 1851 : 199 (226). Lectotype ¢ (by designation of Crosskey, 1971 : 278),
TaAsMANIA (MNHN, Paris) [examined]. — Tasm.

obtusa Walker, 1852 : 274 (Tachina). Lectotype 9 (by fixation of Townsend, 1932 : 40, as
‘Ht’), NEw SoutH Wares (BMNH, London) [examined]. — N.S.W.

stohida Walker, 1858 : 195 (Echinomyia). Holotype gj, New SoutH WaLEs (BMNH, London)

[examined].

ochriventris Malloch, 1929b : 287. Holotype 9, New SoutH Wa tes: Barrington Tops
(ANIC, Canberra) [examined]. —N.S.W., Tasm., Vict.

sinuata Donovan, 1805: plate fig. (unnumbered) & description (unpaginated) (Musca).
Type(s) [3g], AusTRALIA (lost). — A.C.T., N.S.W., QLp, Tasm., Vict., W.A.

Publication date of this name has usually been cited as 1798 in error. Type-material
of sinuata has never been located and is presumed lost (Townsend’s, 1932 : 40, statement
of 2 holotype in London is in error). The identity is certain from Donovan’s coloured
figure, painted it appears from a ¢ specimen.

bura Walker, 1849: 760 (Tachina). Lectotype ¢ (by present designation), TASMANIA
(BMNH, London) [examined].

ignipennis Brauer, 1899 : 510. [Unavailable name first published as a synonym of sinuata
and no later validation; see Crosskey, 1971 : 278.]

latimana Malloch, 1929b : 287. Holotype 9, QUEENSLAND (AM, Sydney) [examined].

skusei Bergroth, 1894: 73. Type(s) 2, QUEENSLAND: Duaringa, Coomooboolaroo (not
located). QLD.
The type-material of this species has not been found in the Zoological Museum, Helsinki
(the most likely depository) and is possibly lost.
violacescens Enderlein, 1937 : 441 (Microtopeza, sic, lapsus). Lectotype ¢ (by present
designation), QUEENSLAND: Herberton (DEI, Eberswalde).—N.S.W., N.T., Qip, W.A.,
Taso. (?).

136 R. W. CROSSKEY

fallax Hardy, 1939 : 35. Holotype 9, QUEENSLAND: Brisbane (not located). Syn. n.
[sinuata Donovan sensu Malloch (misidentification) }
viridescens Paramonov, 1951 : 765. Holotype ¢, AUSTRALIAN CAPITAL TERRITORY: Canberra
(ANIC, Canberra) [examined]. — A.C.T., W.A.

Subfamily GONIINAE Robineau-Desvoidy
GONIDAE Robineau-Desvoidy, 1830 : 74. Type-genus: Gonia Meigen, 1803.

Tribe ACEMYINI Brauer & Bergenstamm

ACEMYIDAE Brauer & Bergenstamm, 1889: 80. Type-genus: Acemya Robineau-Desvoidy,
1830.

Genus CERACIA Rondani

Ceracia Rondani, 1865 : 221. Type-species: Cevacia mucronifera Rondani, 1865, by monotypy.
(ITALY).

Myothyvia Wulp, 1890 : 208. Type-species: Myothyvia majorina Wulp, 1890, by subsequent
designation of Coquillett (I910 : 573). (MEXICO).

armata Malloch, 1930b : 340 (Myothyria). Holotype 9, NEw SoutH WateEs: Kosciusko
(SPHTM, Sydney) [examined]. Comb.n.—A.C.T., N.S.W., Tasm. (Probably = fergusont).
fergusoni Malloch, 19300 : 339 (Myothyria). Holotype 3, WESTERN AUSTRALIA: Eradu
(SPHTM, Sydney) [examined]. Comb.n.-—A.C.T., N.S.W., N.T., Orb, S.A., Vict., W.A.

Tribe NEAERINI Mesnil
NAEREINA [sic] Mesnil, 1956 : 557. Type-genus: Neaeva Robineau-Desvoidy, 1830.

Genus VORIELLA Malloch

Voriella Malloch, 19306 : 335. Type-species: Voriella uniseta Malloch, 1930, by original
designation (as V. vecedens by lapsus: Malloch, 1931 : 298).

uniseta Malloch, 1930b : 335. Holotype g, NEw Soutu Wa tes: Sydney (ANIC, Canberra)
[examined]. — A.C.T., N.S.W., Vict.
vecedens Malloch, 1930b : 335. [Lapsus for uniseta in citation of Voriella type-species; see
Malloch, 1931 : 298.]
? new genera

Undescribed spp. — N.S.W., Qxp, S.A.

Tribe SIPHONINI Rondani
SIPHONAE Rondani, 1845 : 31. Type-genus: Siphona Meigen, 1803.

Genus ACTIA Robineau-Desvoidy

Actia Robineau-Desvoidy, 1830 : 85. Type-species: Actia pilipennis Robineau-Desvoidy, 1830
(junior secondary homonym of pilipennis Fallén, 1810) [=Roeselia lamia Meigen, 1838].
Suspension of ICZN Rules required (see Sabrosky & Arnaud, 1965 : 1061). (EUROPE).

TACHINIDAE OF AUSTRALIA 137

brevis Malloch, 1930b : 309. Holotype g, NEw SoutH WatEs: Sydney (SPHTM, Sydney)
[examined]. — N.S.W.

darwini Malloch, 1929) : 334. Holotype g¢, NorTHERN TERRITORY: Darwin (SPHTM,
Sydney) [examined]. — N.T., QLD.

eucosmae Bezzi, 1926 : 239. Holotype 9, QUEENSLAND: Milton Farm (publ. as ‘Brisbane’)
(BMNH, London) [examined]. —N.S.W., Orp, S.A.

lata Malloch, 1930b : 307. Holotype g, NEw SoutH Wa tes: Sydney (SPHTM, Sydney)
[examined]. — N.S.W.

quadriseta Malloch, 1936 : 20. Holotype 9, NEw SoutH WaLEs: Nyngan (SPHTM, Sydney)
fexamined].—N.S.W.

Genus CEROMYA Robineau-Desvoidy

Ceromya Robineau-Desvoidy, 1830 : 86. Type-species: Cevomya testacea Robineau-Desvoidy,
1830 [=Tachina bicolor Meigen, 1824], by subsequent designation of Coquillett (1910 : 520).
(EUROPE).

Schizoceromyia Townsend, 1926¢ : 542. Type-species: Schizotachina fergusoni Bezzi, 1923, by
original designation. Syn. n.

Schizactiana Curran, 19276 : 356 (as subg. of Actia). Type-species: Actia (Schizactiana) valida
Curran, 1927, by original designation. Syn. n.

Ceromyia. Incorrect subsequent spelling of Cevomya Robineau-Desvoidy.

Schizactina. Incorrect subsequent spelling of Schizactiana Curran (Hardy, 1959 : 213).

fergusoni Bezzi, 1923 : 657 (Schizotachina). Holotype 3, NEw SoutH WaLEs: Sydney (not

located). Comb. n.—N.S.W.
The holotype was stated by Bezzi to be in the Microbiological Laboratory, Sydney,
but has not been located.
fergussont. Incorrect subsequent spelling of feygusont Bezzi (Curran, 1927) : 355-356).

invalida Malloch, 1930b : 305 (Actia). Holotype g, NEw SoutH WaLEs: Sydney (SPHTM,
Sydney) [examined]. — N.S.W.

norma Malloch, 1929a: 116 (Actia). Holotype g, NEw SoutH WateEs: Como (USNM,
Washington) [examined]. —N.S.W., Vict., W.A.

parviseta Malloch, 1930b : 308 (Actia). Holotype g, NEw SoutH WALEs: Sydney (SPHTM,
Sydney) [examined]. — N.S.W.

valida Curran, 19276 : 356 (Actia). Holotype gj, QUEENSLAND: Palmerston (DEI, Eberswalde)
[examined]. Comb. n.—QLp.

Genus PERIBAEA Robineau-Desvoidy

Herbstia Robineau-Desvoidy, 1851 : 184. Type-species: Herbstia tibialis Robineau-Desvoidy,
1851, by monotypy. [Junior homonym of Herbstia Edwards, 1834.] (FRANCE).

Peribaea Robineau-Desvoidy, 1863 : 720. Type-species: Pervibaea apicalis Robineau-Desvoidy,
1863 [=Herbstia tibialis Robineau-Desvoidy, 1851], by subsequent designation of Coquillett
(1910 : 587). (FRANCE).

Strobliomyia Townsend, 1926) : 31. Type-species: Thryptocera fissicornis Strobl, 1910, by
original designation. (AUSTRIA).

Eogymnophthalma Townsend, 1926a : 35. Type-species: Eogymnophthalma orientalisTownsend,
1926 [=Tachina orbata Wiedemann, 1830], by original designation. (SUMATRA).

Talavactia Malloch, 1930b : 305 (as subg. of Actia). Type-species: Actia (Talaractia) baldwini
Malloch, 1930, by original designation.

Tavavactia. Incorrect multiple original spelling of Talavactia Malloch.

Uschizactia Townsend, 1934 : 248. Type-species: Actia uniseta Malloch, 1930, by original
designation. (MAaAtraya).

138 i. W. CROSSKIEY

argentifrons Malloch, 1930) : 309 (Actia). Holotype g, NEw SoutH WateEs: Sydney
(SPHTM, Sydney) [examined]. Comb. n.—N.S.W., OLp.
angustifrons. Incorrect subsequent spelling, lapsus for argentifrons Malloch (Hardy,
1959 : 213).
baldwini Malloch, 1930b : 306 (Actia). Holotype g, QUEENSLAND: Palm Is. (SPHTM,
Sydney) [examined]. Comb. n.— QOLp.
orbata Wiedemann, 1830 : 336(Tachina). Neotype® (by designation of Crosskey, 1967d : 106),
EASTERN Inp1IA: Assam, Azra (BMNH, London) [examined]. Comb. n.—N.S.W., QLp;
New GUINEA; widespread in ORIENTAL REGION, MIDDLE East, AFRICA.
aegyptia Villeneuve, 1912 : 508 (Gymnopareia). Lectotype ¢ (by designation of Crosskey,
1966b : 108), Eeypr: Qaliib (BMNH, London) [examined].
orientalis Townsend, 1926a : 35 (Eogymnophthalma). Lectotype ¢ (by fixation of Townsend,
1940 : 213), SUMATRA: Fort de Kock (ZM, Amsterdam) [examined].
nigritula Malloch, 1930b : 309 (Actia). Holotype 9, QUEENSLAND: Cairns (SPHTM, Sydney)
[examined].
monticola Malloch, 1930c : 143 (Actia). Holotype g [head lost], PHILIppinres: Negros,
Cuernos Mts. (USNM, Washington) [examined].
votundipennis Malloch, 1930c : 143 (Actia). Holotype 9 [head lost], Purrrppines: Negros,
Cuernos Mts. (USNM, Washington) [examined].
sovorcula Mesnil, 1954a:16 (Strobliomyia). Holotype 9, ZairE ReEpusiic: Rutshuru
(MRAC, Tervuren).

plebeia Malloch, 1930b : 310 (Actia). Holotype g, NEw SoutH WaLEs: Coramba (SPHTM,
Sydney) [examined]. Comb. n.-—N.S.W., Qrp.
plebia. Incorrect subsequent spelling of plebeia Malloch (Hardy, 1959 : 213).

Tribe BLONDELIINI Robineau-Desvoidy

BLONDELIDAE Robineau-Desvoidy, 1863 (2): 24. Type-genus: Blondelia Robineau-
Desvoidy, 1830.

Genus ANAGONIA Brauer & Bergenstamm

Anagonia Brauer & Bergenstamm, 1891 : 348 (44). Type-species: Anagonia spylostoides
Brauer & Bergenstamm, 1891 [= Masicera vufifacies Macquart, 1847], by original designa-
tion and monotypy.

Acephana Townsend, 1916c : 153. Type-species: Masicera rubrifrons Macquart, 1847, [= Masi-
ceva vufifacies Macquart, 1847], by original designation.

Opsophana Townsend, 1916c : 153. Type-species: Masicera vufifacies Macquart, 1847, by
original designation.

anguliventris Malloch, 1932b : 273 (Froggattimyia). Holotype 9, AUSTRALIAN CAPITAL
TERRITORY: Canberra (USNM, Washington) [examined]. Comb. n. —A.C.T.

grisea Malloch, 19300 : 333 (Delta). Holotype 9, WESTERN AusTRALIA: Mullewa (ANIC,
Canberra) [examined]. Comb. n.— W.A.

lasiophthalma Malloch, 1934 : 6 (Froggattimyia). Holotype 9, AUSTRALIAN CAPITAL
TERRITORY: Canberra, Black Mt. (ANIC, Canberra) [examined]. Comb. n.—A.C.T.

lateralis Macquart, 1846 : 291 (163) (Masicera). Lectotype g (by designation of Crosskey,
1971 : 274), AUSTRALIA (BMNH, London) [examined]. Comb. n. —N.S.W., QLD.

major Malloch, 1930b : 334 (Delta). Holotype gf, NEw SoutH WatEs: Eccleston, Allyn River
(SPHTM, Sydney) [examined]. Comb. n.—N.S.W., QLD.

Opaca Malloch, 19300 : 334 (Delta). Holotype 9, NEw SoutH Wares: Sydney (SPHTM,
Sydney) [examined]. Comb. n.—N.S.W.

TACHINIDAE OF AUSTRALIA 139

rufifacies Macquart, 1847 : 87 (71) (Masiceva). Holotype g, ‘Tasmania’ (BMNH, London)
[examined]. — A.C.T., Tasm.
vubrifrons Macquart, 1847 : 85 (69) (Masicerva). Holotype 9, TasmManta (BMNH, London)
[examined].
spylosioides Brauer & Bergenstamm, 1891 : 349 (45). Lectotype gd (by designation of
Crosskey 1966b : 108), Tasmania (NM, Vienna) [examined].
scutellata Malloch, 19306 : 334 (Delta). Holotype g, WESTERN AUSTRALIA: Mullewa (SPHTM,
Sydney) [examined]. Comb. n.-—A.C.T., N.S.W., Vicr., W.A.

Genus COMPSILURA Bouché

Compsilura Bouché, 1834 : 58. Type-species: Tachina concinnata Meigen, 1824, by subsequent
designation of Coquillett (t910 : 526). (EUROPE).

concinnata Meigen, 1824 : 412 (Tachina). Holotype 9, CENTRAL Europe [? AUSTRIA or
GERMANY] (? NM, Vienna, coll. von Winthem). — QLD; widespread OLD WorLD; NortTH
AMERICA (introduced & established).

Genus DELTOMYZA Malloch

Delta Malloch, 1930b : 332. Type-species: Delta austvaliensis Malloch, 1930, by original
designation. [Junior homonym of Delta de Saussure, 1855, and Delia Saalmueller, 1891.}

Deltomyza Malloch, 1931 : 298. [Replacement name for Delta Malloch. ]

Mallochiola Strand, 1932 : 195. [Proposed as replacement name for Delta Malloch; junior
homonym of Mallochiola Bergroth, 1925.]

australiensis Malloch, 1930b : 332 (Delta). Holotype g, WESTERN AusTRALIA: Mullewa
(SPHTM, Sydney) [examined]. — W.A.

Genus FROGGATTIMYIA Townsend

Froggattimyia Townsend, 1916c : 155. Type-species: Froggattimyia hivta Townsend, 1916, by
original designation.

Protomeigenia Townsend, 1916c : 156. Type-species: Protomeigenia aurea Townsend, 1916,
by original designation.

aurea Townsend, 1916c : 156 (Protomeigenia). Holotype g, NEw SoutH WatEs: Manilla
(USNM, Washington) [examined]. — N.S.W.

fergusoni Malloch, 1934: 4. Holotype 3, WESTERN AUSTRALIA: Wyalkatchem (SPHTM,
Sydney) [examined]. — W.A.

hirta Townsend, 1916c : 156. Holotype gj, NEw Soutu WALEs: Sydney (publ. as ‘Mittagong’)
(USNM, Washington) [examined]. — N.S.W., OLp.

nicholsoni Malloch, 1934 :5. Holotype ¢, New Soutu Wa tEs: Lindfield (publ. as ‘Sydney’)
(SPHTM, Sydney) [examined]. — N.S.W., OLD.

tillyardi Malloch, 1934: 6. Holotype g, AUSTRALIAN CAPITAL TERRITORY: Blundell’s (ANIC,
Canberra) [examined]. — A.C.T.

wentworthi Malloch, 1934 : 3. Holotype g, NEw SoutH WaLEs: Wentworth Falls (SPHTM,
Sydney) [examined].—N.S.W., Vict. (Possibly = nicholsont)

Genus LECANIPA Rondani

Lecanipa Rondani, 1859: 156. Type-species: Lecanipa patellifera Rondani, 1859 [=Tachina
leucomelas Meigen, 1824], by original designation. (EUROPE).
Lecanipus. Incorrect subsequent spelling of Lecanipa Rondani.

Undescribed sp. (1) — Vict.
Undescribed sp. (2) —S.A.

140 RK. W. CROSSKEY

Genus LIXOPHAGA Townsend

Lixophaga Townsend, 1908 : 86. Type-species: Livophaga parva Townsend, 1908, by original
designation. (NortH AMERICA).

Microceromasia Villeneuve, 1911 : 82. Type-species: Cevomasia sphenophori Villeneuve, I9ItI,
by original designation. (NEw GUINEA).

sphenophori Villeneuve, 1911 : 82. (Cevomasia). Lectotype 9 (by present designation), NEw
GuINEA: Papua [Laloki River near Port Moresby] (ZM, Amsterdam) [examined]. —- NEw
Gur1nEA; Motuccas. Introduced QOrp (? established), Fiy1 (? established), Hawa
(established).

Genus MEDINODEXIA Townsend

Medinodexia Townsend, 1927a: 57. Type-species: Medinodexia fulviventris Townsend, 1927,
by original designation. (SUMATRA).

morgani Hardy, 1934 : 37 (Zostevomyia). Lectotype ¢ (by present designation), NEw SouTH
Wates: Biniguy (NSWDA, Rydalmere) [examined]. Comb. n.—N.S.W., QLD; CEYLON.

Genus MONOLEPTOPHAGA Townsend

Monoleptophaga Baranov, 1938b : 411. Type-species: Monoleptophaga caldwelli Baranov, 1938,
by original designation.

caldwelli Baranov, 1938 : 411. Lectotype g (by designation of Sabrosky & Crosskey,
1969 : 47), QUEENSLAND: Nambour (BMNH, London) [examined]. — QLD.

Genus PAREUPOGONA Townsend

Pareupogona Townsend, 1916c : 157. Type-species: Masiceva oblonga Macquart, 1847, by
original designation.

oblonga Macquart, 1847 : 86 (70) (Masiceva). Holotype gj, Tasmania (BMNH, London)
[examined]. — Tasm.
simplex Macquart, 1847 : 87 (71) (Masiceva). Holotype g, Tasmania (BMNH, London)
[examined].

Genus PAROPSIVORA Malloch

Paropsivora Malloch, 1934: 7. Type-species: Paropsivora grisea Malloch, 1934, by original
designation.

australis Macquart, 1847 : 84 (68) (Degeeria). Holotype 9, Tasmania (BMNH, London)
{fexamined]. Comb. n.—Tasm.

graciliseta Macquart, 1847 : 88 (72) (Phorocerva). Holotype g, TasmMania (BMNH, London)
[examined]. Comb. n.—N.S.W., OLp, Taso.

acutangulata Macquart, 1848 : 208 (48) (Phorocera). Holotype g, Austratia (BMNH,

London) [examined]. Syn. n.

grisea Malloch, 1934: 7. Holotype 9, AUSTRALIAN CAPITAL TERRITORY: Blundell’s (ANIC,
Canberra) [examined) — A.C.T.

tessellata Macquart, 1846 : 293 (165) (Phorocera). Holotype g, Tasmania (MNHN, Paris)
[examined]. Comb. n.— Taso.

TACHINIDAE OF AUSTRALIA I4t

Genus PILIMYIA Malloch

Pilimyia Malloch, 1930b : 329. Type-species: Pilimyia lasiophthalma Malloch, 1930, by
original designation.

lasiophthalma Malloch, 1930) : 329. Holotype g, NEw SoutH WALEs: Blue Mts (SPHTM,
Sydney) [examined]. — N.S.W.

lateralis Macquart, 1846 : 293 (165) (Phorocera). Lectotype ¢ (by designation of Crosskey,
1971 : 282), ‘TAsMANIA’ (MNHN, Paris) [examined]. Comb. n.—Tasm. (?).

Genus TRIGONOSPILA Pokorny

Trigonospila Pokorny, 1886 : 191. Type-species: Trigonospila picta Pokorny, 1886 [=Tachina
ludio Zetterstedt, 1848], by monotypy. (EUROPE).

Zosteromyia Brauer & Bergenstamm, 1891 : 376(72). Type-species: Myobia cingulata Macquart
sensu Brauer & Bergenstamm (misidentification) [=Zostevomyia braueri Townsend,
1933], by original designation. (Note: Townsend’s belief that Brauer & Bergenstamm
misidentified congulata Macquart needs confirmation.)

Zosteromyiopsis Townsend, 1933 : 456. Type-species: Myobia cingulata Macquart, 1851, by
original designation.

braueri Townsend, 1933: 457 (Zosteromyia). Holotype g, Tasmania (NM, Vienna).
Comb. n. — QOxp, Taso.

brevifacies Hardy, 1934 : 36 (Zostevomyia). Lectotype 9 (by present designation), NEw
SoutH WaLEs: Tooloom (QM, Brisbane) [examined]. Comb. n.—N.S.W., Tas.
(Possibly = braueri).

cingulata Macquart, 1851 : 179 (206) (Myobia). Lectotype g (by designation of Crosskey,
1971 : 279), TasmMANIA (MNHN, Paris) [examined].—N.S.W., Qxp, Tasm., Vict.

fasciata Hardy, 1934 : 35 (Zosteromyia). Syntypes 3, 9, Tasmania: Hobart, Mt Wellington
& Victoria: Melbourne (not located). Comb. n. -—Tasm., Vict.

The generic position requires confirmation when type-material located.

Genus ZENARGOMYIA Crosskey
Zenargomyia Crosskey, 1964 :18. Type-species: Zenargomyia moorvet Crosskey, 1964, by
original designation.

moorei Crosskey, 1964 : 20. Holotype g, NEw SoutH WaALEs: Matong S.F. (AM, Sydney)
[examined]. — N.S.W.

Genus ZOSTEROMEIGENIA Townsend

Zosteromeigenia Townsend, 1919a : 579. Type-species: Zosteromeigenita mima Townsend, IgI9Q,
by original designation.

mima Townsend, 1919a: 579. Holotype ¢, QUEENSLAND: Hamilton, Upper North Pine
(USNM, Washington) [examined]. — QLD.
longicornis Hardy, 1934 : 36 (Zosteromyia). Lectotype ¢ (by present designation), QUEENS-
LAND: Mt. Glorious (UQ, Brisbane) [examined]. Syn. n.

Unplaced species of Blondeliini

minor Hardy, 1934 : 36 (Zosteromyia). Syntypes g, TasMANIA: Strahan (not located), —
TasM.

142 KR. We CROSSKEY

similis Macquart, 1851 : 167 (194) (Masicera). Holotype 9, ‘Tasmania’ (MNHN, Paris)
[examined]. — Tasm. (?).
Holotype of this species is in dreadful condition but is apparently a Blondeliine. The
prosternum is haired and the parafacials fully haired.

Tribe EXORISTINI Robineau-Desvoidy
EXORISTIDAE Robineau-Desvoidy, 1863 (1) : 244. Type-genus: Ewvorista Meigen, 1803.

Genus AUSTROPHOROCERA Townsend

Austrophoroceva Townsend, 1916¢ : 157. Type-species: Phoroceva biserialis Macquart, 1847,
by original designation

Glossosalia Mesnil, 1947 : 62 (as subg. of Spoggosia Rondani). [Unavailable: no fixation of a
type-species from two included species. |

Glossosalia Mesnil, 1960 : 606 (as subg. of Spoggosia Rondani). Type-species: Phorocera
gvandis Macquart, 1851, by original designation.

biserialis Macquart, 1847 : 89 (73) (Phorocerva). Lectotype ¢ (by fixation of Townsend,
1940 : 158), TASMANIA (BMNH, London) [examined]. — Tasm., Vict.

grandis Macquart, 1851 : 171 (198) (Phovocerva). Holotype g, AusTRALIA (MNHN, Paris)
[examined]. — N.S.W., S.A., OLD; NEW GUINEA; ORIENTAL REGION.

Undetermined sp. — N.S.W.

Genus CHAETORIA Becker

Chaetoria Becker, 1908 : 113. Type-species: Chaetovia stylata Becker, 1908, by monotypy.
(CANARY ISLANDS).

Vorina Malloch, 1930b : 321. Type-species: Vorina setibasis Malloch, 1930, by original
designation.

setibasis Malloch, 1930b : 321 (Vorina). Holotype 9, NEw SoutH WaLEs: Sydney (SPHTM,
Sydney) [examined]. — N.S.W.

Genus EOZENILLIA Townsend

Eozenillia Townsend, 1926c : 542. Type-species: Eozenillia equatorialis Townsend, 1926, by
original designation. (SINGAPORE).

remota Walker, 1852 : 280 (ITachina). Holotype 9, NEw SoutH WaLEes (BMNH, London)
[examined]. Comb. n.—N.S.W., S.A.

flavipalpis Macquart, 1855 : 122 (102) (Phorocera). Holotype 9 [not ¢], NEw SoutH WALEs:
Sydney (BMNH, London) [examined].

Genus EXORISTA Meigen

Exorista Meigen, 1803 : 280. Type-species: Musca larvarum Linnaeus, 1758, by monotypy.
(EUROPE).

Thrycolyga Rondani, 1856: 68. Type-species: Thrycolyga nova Rondani, 1856, by original
designation. (ITALY).

Eutachina Brauer & Bergenstamm, 1889 : 98 (30). Type-species: Musca larvarum Linnaeus,
1758, by monotypy. (EuRopr). [Junior objective synonym of Evorista.]

Podotachina Brauer & Bergenstamm, 1891 : 350 (46). Type-species: Tachina sorbillans
Wiedemann, 1830, by subsequent designation of Townsend (1916a: 8). (CANARY
IsLANDS).

TACHINIDAE OF AUSTRALIA 143

Tricholyga. Incorrect subsequent spelling of Thrycolyga Rondani.
[Tachina Meigen sensu authors (misidentification) }

auriceps Macquart, 1851 : 158 (185). Holotype g, [TASMANIA: publ. as Oceania] (MNHN,
Paris) [examined].—Tasm. (Probably = flaviceps).

Mesnil (1960 : 578) placed auriceps (with the wrongly cited date 1849) as a synonym
of sorbillans (Wiedemann) and Herting (1962 : 80) wrote that ‘Die Type von sorbillans
Wied. aus Teneriffa . . . ist mit auwviceps Macq. identisch’. Though auriceps undoubtedly
belongs to the sorvbillans complex it is here considered insufficiently substantiated that
auriceps is synonymous with sorbillans s. str. (particularly as the male genitalia of awriceps
holotype have never been examined) and the name auriceps is here maintained as valid
pending further study.

coras Walker, 1849 : 785 (Tachina). Holotype ¢ [abdomen lost], WESTERN AUSTRALIA:
Perth (BMNH, London) [examined]. — N.S.W., W.A.

curriei Curran, 1938: 197 (Thrycolyga). Holotype g, QUEENSLAND: Lawnton (BMNH,
London) [examined]. —- QLp.

doddi Curran, 1938 : 201 (Zenillia). Holotype J, QUEENSLAND: Herberton (DEI, Eberswalde)
fexamined]. — OLD.

flaviceps Macquart, 1847 : 83 (67). Holotype g, Tasmania (BMNH, London) [examined]. —
N-S.W., S.A., Tasm., Vict.

consanguinea Macquart, 1851 : 167 (194) (Masicera). Holotype g, [TasmMania: publ. as

Oceania] (MNHN, Paris) [examined]. Syn. n.

mungomeryi Baranov, 1938) : 410 (Eutachina). Lectotype g (by designation of Sabrosky
& Crosskey, 1969 : 43), QUEENSLAND: Gordonvale (BMNH, London) [examined]. — Orb.

psychidivora Coquillett, 1904 : 137 (Tachina). Holotype gj, WESTERN AusTRALIA (USNM,
Washington) [examined]. — W.A.

sorbillans Wiedemann, 1830 : 311 (Tachina). Lectotype g (by fixation of Townsend, 1932 : 45),
Canary IsLanps: Teneriffe (NM, Vienna).—QLrp; NEw GUINEA; widespread S. PALaeE-
ARCTIC, ORIENTAL, ETHIOPIAN REGIONS.

This species was described from at least two syntypes, as Wiedemann stated ‘In v.
Winthem’s und meiner Sammlung’. There are no surviving specimens in Wiedemann’s
collection, but an original syntype exists in von Winthem’s collection (Vienna). This
specimen was cited by Townsend (1932 : 45) as ‘male Ht [=holotype] in Wien, labelled
“Teneriffa : Coll Winthem’’’, who thus provided a valid fixation of the specimen as
lectotype. The lectotype was not available during the preparation of the present work,
but has been examined by Herting (1962 : 80).

Confirmation is needed that Queensland specimens are truly conspecific with the
lectotype.

tristis Curran, 1938 : 203 (Zenillia). Holotype 9, QUEENSLAND: Cairns (DEI, Eberswalde)
[examined]. — OLp.

Genus HILLOMYIA Crosskey nom. n.

Hillia Malloch, 19296 : 328. Type-species: Hillia polita Malloch, 1929, by original designation.
[Junior homonym of Hillia Grote, 1883.|

Hillomyia Crosskey nom.n. [New name for Hillia Malloch.]

polita Malloch, 1929b : 328. Holotype 9, NoRTHERN TERRITORY: Darwin (ANIC, Canberra)
[examined]. Comb. n.—N.T.

Genus SPOGGOSIA Rondani*

Spoggosia Rondani, 1859: 182. Type-species: Spoggosia occlusa Rondani, 1859 [=Salia
echinura Robineau-Desvoidy, 1830], by monotypy. (EUROPE).
* See Appendix, p. 209.

144 R. W. CROSSKEY

micropalpis Malloch, 19306 : 321 (Stomatomyia). Holotype 3, NEw SoutH WaLEs: Sydney
(SPHTM, Sydney) [examined]. — A.C.T., N.S.W., Tasm., Vict.
Undescribed sp. — N.S.W.

Genus STOMATOMYIA Brauer & Bergenstamm

Stomatomyia Brauer & Bergenstamm, 1889: 98 (30). Type-species: Chetogena filipalpis
Rondani, 1859, by monotypy. (ITALy).

Plagiprospherysa Townsend, 1892a: 113. Type-species: Plagiprospherysa valida Townsend,
1892 [=Prospherysa parvipalpis Wulp, 1890], by original designation. (NORTH AMERICA).

acuminata Rondani, 1859 : 180 (Chetogena). Syntypes [? sex], Iraty (MZ, Florence). — N.S.W.
Qtp, W.A. (vide Malloch, 1930b : 320); S. PALAEARCTIC REG., JAPAN; ORIENTAL REG.
including INDONESIA.

tricholygoides Bezzi, 1928 : 205. Holotype g¢, Fij1: Ovalau (BMNH, London) [examined]. —
N.S.W., OLD (?); Fiy1, NEw GUINEA, SOLOMONS.

Tribe ETHILLINI Mesnil
ETHYLLINA [sic] Mesnil, 1944 : 23. Type-genus: Ethilla Robineau-Desvoidy, 1863.

Genus ETHILLA Robineau-Desvoidy

Ethilla Robineau-Desvoidy, 1863 : 202. Type-species: Tachina aemula Meigen, 1824, by
original designation. (EUROPE).

Ethylla. Incorrect subsequent spelling of Ethilla Robineau-Desvoidy.

translucens Macquart, 1851 : 162 (189) (Evorista). Holotype g, Tasmania (MNHN, Paris)
[examined]. Comb. n.-—S.A., Tasm., Vict.

Genus PHOROCEROSOMA Townsend

Phorocevosoma Townsend, 1927a: 61. Type-species: Phovocerosoma forte Townsend, 1927
[=Masicera vicaria Walker, 1847], by original designation. (SUMATRA).

cilipes Macquart, 1847 : 88 (72) (Phoroceva). Holotype g, Tasmania (BMNH, London)
[examined]. Comb. n.— Tasm.
vufomaculata Macquart, 1851: 160 (187) (Evorista). Holotype g, Tasmania (MNHN,
Paris) [examined]. Syn. n.
This species is certainly an ethilline, but there are some differences from Phorocerosoma
and inclusion in this genus is an interim measure.
postulans Walker, 1861a : 240 (Nemoraea). Holotype ¢ [head lost], NEw Guinea: Dorey
(BMNH, London) [examined].—N.T., Q1rp; NEw GuINEA; SoLomons; widespread
ORIENTAL REGION and tropical AFRICA.
mysolana Walker, 1864 : 213 (Masiceva). Holotype 9, InponEs1a: Moluccas, Miso6l (publ.
as Mysol) (BMNH, London) [examined].
anomala Baranov, 1936: 99. Lectotype 9 (by designation of Crosskey, 1966b : 108),
Formosa: Koshun, Kankau (DEI, Eberswalde) [examined].
nitidicauda Curran, 1938 : 202 (Zenillia). Holotype 3, QUEENSLAND: Cairns (SPHTM,
Sydney) [examined].

TACHINIDAE OF AUSTRALIA 145

Genus MYCTEROMYIELLA Mesnil

Mycteromyia Mesnil, 1950a : 107. Type-species: Mycteromyia laetifica Mesnil, 1950, by original
designation. (New Guinea). [Junior homonym of Mycteromyia Philippi, 1865.]
Mycteromyiella Mesnil, 1965 : 232. [Replacement name for Mycteromyia Mesnil.]

Undescribed sp. — N.S.W.

Tribe WINTHEMIINI Townsend
WINTHEMITAE Townsend, 1913 : 52. Type-genus: Winthemia Robineau-Desvoidy, 1830.

Genus CRYPSINA Brauer & Bergenstamm

Crypsina Brauer & Bergenstamm, 1889 : 97 (29). Type-species: Crypsina prima Brauer &
Bergenstamm, 1889, by original designation and monotypy.

Amplipila Curran, 1927c : 446. Type-species: Amplipila versicoloy Curran, 1927 [=Crypsina
prima Brauer & Bergenstamm, 1889], by original designation.

prima Brauer & Bergenstamm, 1889 : 97 (29). Holotype 9, QUEENSLAND: Rockhampton
(NM, Vienna) [examined]. — OLD.

versicoloy Curran, 1927¢ : 446 (Amplipila). Holotype g, QUEENSLAND: Herberton (DEI,

Eberswalde) [examined].

Genus NEMORILLA Rondani

Nemorilla Rondani, 1856: 66. Type-species: Tachina maculosa Meigen, 1824, by original
designation. (EUROPE).

Undetermined sp.—Qtp. (The mis-associated 9 paralectotype of Sisyropa cinerea Brauer &
Bergenstamm from Cape York is a specimen of Nemorilla but the species has not been
determined).

Genus WINTHEMIA Robineau-Desvoidy

Winthenna Robineau-Desvoidy, 1830 : 173. Type-species: Tachina variegata Meigen, 1824, by
subsequent designation of Robineau-Desvoidy (1863 : 207). (EUROPE).

Pseudokea Townsend, 1928 : 393. Type-species: Pseudokea neowinthemioides Townsend, 1928,
by original designation. (PHILIPPINEs).

Winthemya. Incorrect subsequent spelling of Winthemia Robineau-Desvoidy.

lateralis Macquart, 1843 : 215 (58) (Eurigaster). Holotype g, AustraLtia (MNHN, Paris)
[examined]. — N.S.W., S.A., Tasm.
brevisetosa Macquart, 1846 : 282 (154) (Nemoraea). Holotype g, Tasmania (MNHN, Paris)
[examined]. Syn. n.
lata Macquart, 1848 : 207 (47) (Evorista). Holotype g, AusTRaLtiA (BMNH, London)
[examined]. Syn. n.
marginata Macquart, 1851 : 161 (188) (Evorista). Lectotype § (by designation of Crosskey,
1971 : 269), TasmMANIA (MNHN, Paris) [examined]. Syn. n.
albiceps Malloch, 1930b : 349. Holotype g, NEw Sout WALEs: Sydney (SPHTM, Sydney)
[examined]. Syn. n.
albicens. Incorrect multiple original spelling of albiceps Malloch.
neowinthemioides Townsend, 1928 : 394 (Pseudokea). Holotype g, PHILIPPINES: Mindanao,
Cagayan (USNM, Washington) [examined].—N.S.W., QLD; NEw GUINEA, PHILIPPINES,
INDONESIA.
diversa Malloch, 1930b : 348. Holotype g, New Soutu WateEs: Killara, Allowrie (SPHTM,
Sydney) [examined]. Syn. n.
K

146 R. W. CROSSKEY

trichopareia Schiner, 1868 : 327 (Exorista). Type(s) [? sex], [AusTRaLt1a] (not located,
possibly lost).- QLp. (Confirmation of identity required).

This species was, as Malloch (19300 : 349) stated, unsatisfactorily described by Schiner
who did not state the number of specimens, or their sex or their locality of origin. Nothing
in the description indicates positively that Winthemia is the correct genus, and Malloch’s
(1930b) placement was made by guesswork from Brauer & Bergenstamm’s (1891 : 441)
earlier placement in Chaetolyga; it is from their statement of ‘Australien’ that the proven-
ance of Schiner’s species is known. The original type-material has never been studied
and cannot now be found amongst Schiner’s types in NM, Vienna.

Tribe CARCELIINI Townsend
CARCELIIAE Townsend, 1913 : 52. Type-genus: Carcelia Robineau-Desvoidy, 1830.

Genus ARGYROPHYLAX Brauer & Bergenstamm

Argyvophylax Brauer & Bergenstamm, 1889 : 163 (95). Type-species: Tachina albincisa
Wiedemann, 1830, by original designation and monotypy. (West INDIEs).

solomonica Baranov, 1938a : 170 (Bactromyia, as subsp. of fyansseni Baranov). Lectotype ¢
(by designation of Sabrosky & Crosskey, 1969 : 36), SOLOMON IsLanDs: Russell Is. (BMNH,
London) [examined]. — QLD; NEw GUINEA, SOLOMONS.

Genus ARGYROTHELAIRA Townsend

Argyvothelaiva Townsend, 1916a : 311. Type-species: Argyrothelaiva froggattii1 Townsend, 1916,
by original designation. (SOLOMON ISLANDs).

melancholica Mesnil, 1944 : 29 (Carcelia). Holotype 9 [head and most legs lost], QUEENS-
LAND: Cairns (DEI, Eberswalde) [examined]. Comb. n.—QO1Lp; NEw GUINEA.

Genus CARCELIA Robineau-Desvoidy

Caycelia Robineau-Desvoidy, 1830 : 176. Type-species: Carcelia bombylans Robineau-Desvoidy,
1830, by subsequent designation of Townsend (1916a : 6). (EUROPE).

Subgenus CARCELIA Robineau-Desvoidy

Carcelia Robineau-Desvoidy, 1830 : 176. Type-species: Carcelia bombylans Robineau-Desvoidy,
1830, by subsequent designation of Townsend (1916a : 6). (EUROPE).

hardyi Curran, 1938 : 200 (Zenillia). Holotype 3, QUEENSLAND: Brisbane (SPHTM, Sydney)
[examined]. — QLp, Vict. (?).
tasmanica Robineau-Desvoidy, 1863 : 240. Holotype g, TAsMANIA (lost). — Tas.
scutellavis Robineau-Desvoidy, 1863 : 240 (Phoroceva). [Unavailable name, first published
as a Synonym. |

Robineau-Desvoidy named C. tasmanica for a male specimen said by him to be in the
Paris Museum and to be labelled ‘Phorocera scutellaris’ by Macquart. Macquart did
not publish this name, but it was cited by Robineau-Desvoidy in synonymy with tasmanica.
The name scutellavis is therefore attributable to Robineau-Desvoidy and is unavailable
under Article 11(d) of the ICZN. The type-specimen cannot be found in MNHN, Paris,
and is presumed lost.

TACHINIDAE OF AUSTRALIA

Subgenus SENOMETOPIA Macquart

Senometopia Macquart, 1834 : 296 (as genus). Type-species: Carcelia aurifrons Robineau-
Desvoidy, 1830 [=Tachina excisa Fallén, 1820], by subsequent designation of Townsend
(1916a : 8). (EUROPE).

Stenometopia Agassiz, 1846 : 351. Unjustified emendation of Senometopia Macquart.

Eocarcelia Townsend, 1919a : 582. Type-species: Eocarcelia ceylanica Townsend, 1919, by
original designation. (CEYLON). Syn. n.

Eocarceliopsis Townsend, 1928 : 392. Type-species: Eocarceliopsis bakeri Townsend, 1928, by
original designation. (PHILIPPINES). Syn. n.

Eucarcelia Baranov, 1934) : 393. Type-species: Tachina excisa Fallén, 1820, by original
designation. (EUROPE).

Dicephalomyia Malloch, 1935 : 337. Type-species: Dicephalomyia rufiventris Malloch, 1935,
by original designation. (BORNEO). Syn. n.

cinerea Brauer & Bergenstamm, 1891 : 346 (42) (Sisyropa). Lectotype ¢ (by present designa-
tion), QUEENSLAND: Rockhampton (NM, Vienna) [examined]. —- QLD; NEw GUINEA.

cosmophilae Curran, 1938 : 200 (Zenillia). Holotype g, QUEENSLAND (BMNH, London)
[examined]. —- QLD.

murina Curran, 1938 : 198 (Zenilia). Holotype g, NEw SoutH WaLEs: Wee Waa (BMNH,
London) [examined]. — N.S.W., OQLp.

noctuae Curran, 1938 : 199 (Zenillia). Holotype J, QUEENSLAND: Lawnton (BMNH, London)
[examined].—Qtp. (Possibly = zlota Curran, 1927, of Ethiopian Region).

Genus CARCELIMYIA Mesnil

Carcelimyia Mesnil, 1944 : 26. Type-species: Evorista dispay Macquart, 1851, by original
designation.

dispar Macquart, 1851 : 159 (186) (Evorista). Lectotype 3 (by designation of Crosskey,
1971 : 268), AUSTRALIA (MNHN, Paris) [examined).— N.S.W., N.T., W.A.

Tribe ANACAMPTOMYIINI Townsend

ANACAMPTOMYITINI Townsend, 1936 : 35, 38, 41. Type-genus: Anacamptomyia Bischof,
1904.

Genus ANACAMPTOMYIA Bischof

Anacamptomyia Bischof, 1904 : 79. Type-species: Anacamptomyia africana Bischof, 1904, by
monotypy. (SoutTH AFRICA).

Vespivora Malloch, 1930) : 347. Type-species: Vespivova nigviventris Malloch, 1930, by
original designation.

nigriventris Malloch, 19306 : 347 (Vespivora). Holotype 3, QUEENSLAND: Eidsvold (ANIC,
Canberra) [examined]. — N.S.W., QLD.

Genus EUVESPIVORA Baranov

Ewvespivova Baranoy, 1942: 161. Type-species: Euvespivora orientalis Baranov, 1942, by
original designation. (JAVA).

Xenostuymia Mesnil, 1944 : 26. Type-species: Xenosturmia testaceipes Mesnil, 1944 [=Eury-
gastey decipiens Walker, 1859], by original designation. (NEw Britain).

148 R. W. CROSSKEY

decipiens Walker, 1859 : 100 (Eurygaster). Holotype 9, Aru Istanps (BMNH, London)
[examined] — N.S.W., OLp; Maraya, Aru IsLANDS, SOLOMONS, NEw CALEDONIA.
salomonica Baranov, 1942 : 163. Holotype 9, Sotomon IsLanps: Tulagi (BMNH, London)
[examined].
testaceipes Mesnil, 1944: 26 (Xenosturmia). Holotype 9, NEw Britain: Kinigunang
(DEI, Eberswalde) [examined].

Genus KORALLIOMYIA Mesnil

Kovalliiomyia Mesnil, 1950a: 114. Type-species: Kovalliiomyia portentosa Mesnil, 1950, by
original designation. (INDIA).

Undetermined spp. (possibly portentosa Mesnil). — QLD.

Tribe STURMIINI Robineau-Desvoidy

STURMIDAE Robineau-Desvoidy, 1863 (1) : 885. Type-genus: Sturmia Robineau-Desvoidy,
1830.

Genus ANAMASTAX Brauer & Bergenstamm

Anamastax Brauer & Bergenstamm, I891 : 349 (45). Type-species: Blepharipeza goniaeformis
Macquart sensu Brauer & Bergenstamm (misidentification) [= Anamastax australis Town-
send, 1933], by original designation.

braueri Hardy, 1938 : 62 (ITvitaxys, as replacement name for austvalis Townsend). — N.S.W.,

QLD.
australis Townsend, 1933 : 473. Holotypeg, QUEENSLAND (NM, Vienna).

Hardy (1938 : 59 treated Anamastax as a synonym of Tritaxys, thereby bringing A. australis
Townsend into the same genus as T. australis Macquart. He therefore published the
replacement name brauevi for the secondarily homonymous australis Townsend.
Regrettably, as this change was made prior to 1960, the name braueri stands valid for the
species under the present ICZN even though the species involved are no longer considered
congeneric.

Undescribed sp. — N.S.W.

Genus ARRHENOMYZA Malloch

Arrhenomyza Malloch, 19296 : 322. Type-species: Avrhenomyza conspicua Malloch, 1929, by
original designation.

conspicua Malloch, 19296 : 322. Holotype J, WESTERN AUSTRALIA: Eradu, near Geraldton
(ANIC, Canberra) [examined]. — W.A.

Genus BLEPHARELLA Macquart

Blepharella Macquart, 1851 : 176 (203). Type-species: Blepharella latevalis Macquart, 1851,
by monotypy. (INp1A).

Podomyia Brauer & Bergenstamm, 1889 : 96 (28). Type-species: Eurigaster setosa Doleschall,
1858 [=Blepharella lateralis Macquart, 1851], by original designation. (AMBOYNA).

Phryxosturmia Townsend, 19274 : 68. Type-species: Phryxosturmia jacobsont Townsend, 1927
[=Blepharella lateralis Macquart, 1851], by original designation. (SUMATRA).

Apilia Malloch, 1930b : 345. Type-species: Apilia cilifera Malloch, 1930 [=Blepharella
lateralis Macquart, 1851], by original designation.

—

TACHINIDAE OF AUSTRALIA 149

lateralis Macquart, 1851 : 177 (204). Holotype g, Inp1a: Pondicherry (MNHN, Paris)
[examined].—- QLrp; NEw GUINEA, SOLOMONS, widespread ORIENTAL REGION.
maculata Macquart, 1851 : 173 (200) (Phorocera). Holotype g, AustRaLiIa (MNHN, Paris)
fexamined]. Syn. n.
cilifera Malloch, 1930b : 345 (Apilia). Holotype J, QUEENSLAND: Eidsvold (ANIC, Canberra)
[examined].
(For other synonyms, based on Oriental types, see Crosskey, 1966b : 106)

Genus BLEPHARIPA Rondani

Blepharipa Rondani, 1856 : 71. Type-species: Senometopia ciliata Macquart, 1835 [= Nemoraea
scutellata Robineau-Desvoidy, 1830], by original designation. (FRANCE).

Verveauxia Robineau-Desvoidy, 1863 : 893. Type-species: Verreauxia auripilis Robineau-
Desvoidy, 1863, by original designation. [Junior homonym of Verreauxia Hartlaub,
1856.]

Ugimyia Rondani, 1870 : 137. Type-species: Ugimyia sericarviae Rondani, 1870, by monotypy.
(JAPAN).

Blepharvipoda Brauer & Bergenstamm, 1889: 96 (28). Type-species: Nemovaea scutellata
Robineau-Desvoidy, 1830, by monotypy. (FRANCE). [Junior homonym of Blepharipoda
Randall, 1840].

Crossocosmia Mik, 1890 : 313. Type-species: Ugimyia sericariae Rondani, 1870 (as sericariae
Cornalia), by original designation. [Junior objective synonym of Ugimyia Rondani.]
(JAPAN).

Eopavachaeta Townsend, 1927a: 70. Type-species: Eoparachaeta orientalis Townsend, 1927
[=Tachina zebina Walker, 1849], by original designation. (SUMATRA).

Sumatrosturmia Townsend, 1927a: 70. Type-species: Sumatrosturmia orbitalis Townsend,
1927, by original designation. (SUMATRA).

Indosturmia Townsend, 1932 : 49. Type-species: Crossocosmia indica Brauer & Bergenstamm,
1893, by original designation. (INDIA).

Chrysopygia Townsend, 1933 : 471. Type-species: Chrysopygia auricaudata Townsend, 1933,
by original designation. (JAVA).

auripilis Robineau-Desvoidy, 1863 : 894 (Verreauxia). Holotype g, TAsMANIA (lost).
Comb. n. — Tasm., N.S.W. (?).

coesiofasciata Macquart, 1851 : 165 (192) (Masiceva). Holotype 9, AusTRaLIa (MNHN,
Paris) [examined]. Comb. n.-—N.S.W.

fulviventris Macquart, 1851 : 165 (192) (Masicera). lectotype ¢ (by designation of Crosskey,
1971 : 274), ‘TASMANIA’ (MNHN, Paris) [examined]. Comb. n.—N.S.W., QLp, Taso.
(?). (Possibly = sugens Wiedemann of Oriental Region)

austvalis Walker, 1852 : 279 (Tachina). Holotype g, NEw SoutH WaLEs (BMNH, London)

[examined]. Syn. n.

Genus CALOZENILLIA Townsend

Calozenillia Townsend, 1927a : 67. Type-species: Calozenillia auronigya Townsend, 1927, by
original designation. (SUMATRA).

olmus Walker, 1849 : 775 (Tachina). Holotype 9, AustRaLIa (BMNH, London) [examined]. —

QLD.
picta Curran, 1938 : 202 (Zenillia). Holotype 3, QUEENSLAND: Herberton (DEI, Eberswalde)

[examined]. — QLp.

Genus EURYGASTROPSIS Townsend

Eurygastropsis Townsend, 1916c : 158. Type-species: Eurigaster tasmaniae Walker, 1858, by
original designation.

150 RR, Ws CROSSE ¥

Calopygidia Malloch, 1930b : 349. Type-species: Calopygidia analis Malloch, 1930 [=Eurigaster
tasmaniae Walker, 1858], by original designation.

tasmaniae Walker, 1858 : 197 (Eurigaster). Holotype ¢ [not 9], Tasmania (BMNH, London)
[examined]. — N.S.W., QOLp, Tasm., W.A.; NEw GUINEA (?).
analis Malloch, 19300 : 350 (Calopygidia). Holotype g, NEw Soutu WatEs: Barrington
Tops (ANIC, Canberra) [examined].

Genus PALEXORISTA Townsend

Palexorista Townsend, 1921 : 134. Type-species: Tachina succini Giebel, 1862 [=Masicera
solennis Walker, 1859], by original designation. (Presumed ORIENTAL: succini based on
holotype in copal, see Crosskey, 1966c : 133).

Sumatrodoria Townsend, 1927a : 64. Type-species: Sumatrodoria summaria Townsend, 1927,
by original designation. (SUMATRA).

Prosturmia Townsend, 1927a : 69. Type-species: Prosturmia profana Townsend, 1927 [= Masi-
ceva solennis Walker, 1859), by original designation. (SUMATRA).

bancrofti Crosskey, 1967c : 85. Holotype g, QUEENSLAND: Burpengary (ANIC, Canberra)
[examined]. — QLD.
curvipalpis Wulp, 1893 : 162 (Crossocosmia). Lectotype ¢ (by designation of Crosskey,
1967c : 68), JavA (RMNH, Leiden) [examined].—Qip; NEw GuINnEA, SoLomons, wide-
spread ORIENTAL REGION.
unisetosa Baranov, 1932 : 75 (Sturmia). Lectotype ¢ (by designation of Crosskey, 1967c : 68),
Formosa: Kankau, Koshun (DEI, Eberswalde) [examined].
lucagus Walker, 1849 : 678 (Tachina). Holotype g, Cuina: Foo-chow-foo (BMNH, London)
[examined]. — N.T.; NEw Guinea; widespread ORIENTAL REGION.
macquarti Crosskey nom. n. [Replacement name for Masicera auriceps Macquart, 1851.]
auriceps Macquart, 1851 : 168 (195) (Masicera). Lectotype 2 (by designation of Crosskey,
1971 : 273), ‘TasManta’ (MNHN, Paris) [examined]. [Junior primary homonym of
Masicera auriceps Macquart, 1843.]
solennis Walker, 1859 : 98 (Masiceva). Holotype ¢ [not 9], Aru IsLanps (BMNH, London)
[examined]. — QLD; widespread ORIENTAL REGION, MELANESIA and MICRONESIA; TONGA.
profana Townsend, 1927a : 69 (Prosturmia). Lectotype g (by designation of Crosskey,
1969 : 100), SUMATRA: Fort de Kock (ZM, Amsterdam) [examined].
inconspicuella Baranov, 1932: 79 (Sturmia). Lectotype g (by designation of Crosskey,
1967c : 57), Formosa: Kankau, Koshun (DEI, Eberswalde) [examined].
(For further synonymy see Crosskey, 1967¢ : 57.)
sororcula Mesnil, 1949b : 30 (Drino). Holotype g, QUEENSLAND: Herberton (DEI, Ebers-
walde) [examined]. — QLp.
subanajama Townsend, 19274 : 69 (Prosturmia). Lectotype 3 (by designation of Crosskey,
1967¢ : 55), SUMATRA: Suban Ajam (ZM, Amsterdam) [examined]. —- QLD; NEw GUINEA,
SoLomons, MaraysiA, INDONESIA.

Genus PALIA Curran

Palia Curran, 1927¢ : 443. Type-species: Palia aureocauda Curran, 1927, by original designation.

aureocauda Curran, 1927¢ : 444. Holotype g, QUEENSLAND: Kuranda (DEI, Eberswalde)
[examined]. — OLD.

Genus PALIANA Curran

Paliana Curran, 1927¢ : 445. Type-species: Paliana basalis Curran, 1927, by original
designation.

TACHINIDAE OF AUSTRALIA I51

basalis Curran, 1927¢: 445. Holotype g, QuEENSLAND: Kuranda (DEI, Eberswalde)
{examined]. — OLp.

intensa Curran, 1927c¢: 446. Holotype 9, QuEENSLAND: Kuranda (DEI, Eberswalde)
{examined].—~Qzp. (Probably = basalis)

Genus PARADRINO Mesnil

Pavradrino Mesnil, 1949b : 8, 35 (as subg. of Dvino Robineau-Desvoidy). Type-species: Sturmia
halli Curran, 1939, by monotypy. (AFRICA).

laevicula Mesnil, 1951 : 197 (Drino). Holotype 9, Formosa: Kankau, Koshun (DEI, Ebers-
walde) [examined]. —- QLp; NEw GuINEA; widespread ORIENTAL REGION.

Genus PARAGONIA Mesnil

Paragonia Mesnil, 1950a : 106. Type-species: Pavagonia portentosa Mesnil, 1950, by origina
designation.

portentosa Mesnil, 1950a : 106. Holotype 3, WESTERN AUSTRALIA: Waroona (CNC, Ottawa,
ex coll. Mesnil) [examined]. — W.A.

Genus POLYCHAETA Macquart

Polychaeta Macquart, 1851 : 154 (181). Type-species: Polychaeta nigra Macquart, 1851, by
monotypy.
nigra Macquart, 1851 : 154 (181). Holotype 9, ‘Tasmania’ (MNHN, Paris) [examined]. —
A.C.T., N.S.W., Tasm. (?).
subpubescens Macquart, 1851 : 172 (199) (Phoroceva). Holotype 9 [not g], ‘Tasmanta’
(MNHN, Paris) [examined]. Syn. n.

Genus QUADRA Malloch

Quadra Malloch, 1929b : 320. Type-species: Quadva ornata Malloch, 1929, by original designa-
tion.

ornata Malloch, 1929) : 320. Holotype 9, WrEsTERN AUSTRALIA: King George’s Sound
(AM, Sydney) [examined]. — W.A.

Genus SISYROPA Brauer & Bergenstamm

Sisyvopa Brauer & Bergenstamm, 1889 : 163 (95). Type-species: Tachina thermophila Wiede-
mann, 1830, by original designation. (JAVA).

taylori Curran, 1938 : 204 (Zenillia). Holotype 3, QUEENSLAND: Innisfail (SPHTM, Sydney)
[examined]. — N.T., QLp; NEw Guinea, MoLuccas.

Undetermined spp. (with bare eyes). — QLD.

Genus STURMIA Robineau-Desvoidy

Sturmia Robineau-Desvoidy, 1830 : 171. Type-species: Sturmia vanessae Robineau-Desvoidy,
1830 [=Tachina bella Meigen, 1824], by subsequent designation of Robineau-Desvoidy
(1863 : 888).

152 R. W. CROSSKEY

convergens Wiedemann, 1824 : 43 (Tachina). Lectotype 2 (by designation of Crosskey,
1963 : 78), InpD1A (UZM, Copenhagen) [examined].—N.S.W., Qtp; NEw GuINEA; wide-
spread ORIENTAL REGION.
setilatera Wiedemann, 1830 : 321 (Zachina). Lectotype g (by designation of Crosskey,
1966a : 678), ‘IND. OR.’ (probably = Inp1a) (UZM, Copenhagen) [examined].
Undetermined sp. — S.A.

Genus TASMANIOMYIA Townsend

Chlovogastey Macquart, 1851 : 157 (184). Type-species: Chlorogaster tasmanensis Macquart,
1851, by monotypy. [Junior homonym of Chlorogastey Swainson, 1839.]

Tasmaniomyia Townsend, 1916c¢ : 152. . Type-species: Masiceva viridiventris Macquart, 1847,
by original designation.

Chlorogastvina Crosskey, 1967 : 9. [Replacement name for Chlovogastey Macquart.] Syn. n.

tasmanensis Macquart, 1851 : 157 (184) (Chlorogaster). Lectotype g (by designation of
Crosskey, 1971 : 264), ‘TASMANIA’ (MNHN, Paris) [examined]. Comb. n.—Tasm. (?).

viridiventris Macquart, 1847 : 84 (68) (Masicera). Lectotype ¢ (by fixation of Townsend,
1939 : 15), TasmMANIA (BMNH, London) [examined]. — Tasm.*

Genus TRITAXYS Macquart

Tritaxys Macquart, 1847 : 81 (65). Type-species: Tvitaxys austvalis Macquart, 1847, by
monotypy.

Goniophana Brauer & Bergenstamm, 1889 : 97 (29). Type-species: Gonia heterocera Macquart,
1846, by original designation.

Gonanamastax Townsend, 1933 : 472. Type-species: Blepharipeza goniaeformis Macquart, 1846,
by original designation.

Goniophania. Incorrect subsequent spelling of Goniophana Brauer & Bergenstamm (Malloch,
1929a : 113, attributed to Townsend in error).

australis Macquart, 1847 : 82 (66). Lectotype ¢ (by designation of Crosskey, 1971 : 292),
Tasmania (BMNH, London) [examined]. — Tasm.

goniaeformis Macquart, 1846: 285 (157) (Blepharipeza). Lectotype Q (by fixation of
Townsend, 1932: 50), TASMANIA (MNHN, Paris) [examined].—Tasm. (Possibly =
hetevocera).

heterocera Macquart, 1846 : 281 (153) (Gonia). Lectotype g (by designation of Crosskey,
1971 : 269), ‘TAsMANIA’ (MNHN, Paris) [examined]. — N.S.W., OLD, Vict., W.A., TAs. (?).

milias Walker, 1849 : 799 (Gonia). Holotype 9, NEw SoutH WaLeEs: Hunter River (BMNH,
London) [examined]. — N.S.W., OLD.

milas. Incorrect subsequent spelling of milias Walker (Hardy, 1938 : 61).

scutellata Macquart, 1846 : 293 (165) (Phoroceva). Holotype g, Tasmania (MNHN, Paris)

[examined]. Comb. n. — Tas.

Genus UGIMEIGENIA Townsend
Ugimeigenia Townsend, 1916d : 316. Type-species: Ugimeigenia elznevi Townsend, 1916, by
original designation.

elzneri Townsend, 1916d : 316. Holotype g, QUEENSLAND: Banks Island (USNM, Washington)
[examined]. — OLD.

* See Appendix, p. 209.

TACHINIDAE OF AUSTRALIA 153

Genus WINTHELLIA Crosskey

Thyellina Mesnil, 1949a : 70. Type-species: Thyellina brevicornis Mesnil, 1949, by monotypy.
[Junior homonym of Thyellina Agassiz, 1838.]
Winthellia Crosskey, 1967a : 31. [Replacement name for Thyellina Mesnil.]

brevicornis Mesnil, 1949a: 70 (Thyellina). WHolotype 9, QUEENSLAND: Herberton (CNC,
Ottawa, ex coll. Mesnil) [examined]. — QLp.

Genus ZEBROMYIA Malloch

Zebromyia Malloch, 1929 : 321. Type-species: Zebromyia obesa Malloch, 1929 [=Phorocera
ornata Macquart, 1851], by original designation.

ornata Macquart, 1851 : 172 (199) (Phorocera). Lectotype ¢ (by designation of Crosskey,
1971 : 283), TasMANIA (MNHN, Paris) [examined]. — N.S.W., Tasm.
obesa Malloch, 1929 : 321. Holotype g, Tasmanta (AM, Sydney) [examined].

Genus ZYGOBOTHRIA Mik

Zygobothria Mik, 1891 : 193. Type-species: Sturmia atropivora Robineau-Desvoidy, 1830, by
original designation. (EUROPE).

ciliata Wulp, 1881 : 38 (Meigenia). Lectotype ¢ (by designation of Crosskey, 1967d : 104),
SuMATRA: Alahan pandjang (RMNH, Leiden) [examined].— W.A.; NEw GuINEA; wide-
spread ORIENTAL and ETHIOPIAN REGIONS.

atropivora Robineau-Desvoidy, 1830:171 (Stwrmia). Syntypes [¢ 9}, [France] (lost). —
N.S.W., QLD; widespread in OLD WorRLD.

Robineau-Desvoidy based this species on over eighty specimens that Serville reared
from a chrysalis of Sphinx atropos. Both sexes are presumed to have been present in this
number. None of the syntypes have been located or recognized since and all are presumed
lost. (None exist in the remnants of Robineau-Desvoidy’s collection in MNHN, Paris.)

Undetermined sp. — A.C.T.

Unplaced species of Sturmiini

castanea Hardy, 1938 : 63 (Calopygidia). Holotype g, Tasmania: Mt Wellington (not
located). — Tasm.
dissimilis Malloch, 193cb : 343 (Quadra). Holotype g, NEw SoutH WateEs: Como (ANIC,
Canberra) [examined]. — A.C.T., N.S.W.
This species is considered not to be congeneric with ovnata Malloch, type-species of
Quadra, but no generic assignment is possible at present.
Undescribed spp. (? genera) — various states.

Tribe GONIINI Robineau-Desvoidy
GONIDAE Robineau-Desvoidy, 1830 : 74. Type-genus: Gonia Meigen, 1803.

Genus GONIOPHTHALMUS Villeneuve

Goniophthalmus Villeneuve, 1910: 145. Type-species: Goniophthalmus simonyi Villeneuve,
1910, by monotypy. (SOKOTRa).
australis Baranov, 1938) : 405 (Dolichocolon australe). Lectotype g (by designation of

Sabrosky & Crosskey, 1969 : 40), QUEENSLAND: Gympie (BMNH, London) [examined]. —
N.S.W., Oup, W.A. (?).

154 RW: CROSSKEY

rufescens Baranov, 1938b : 406 (Dolichocolon). Lectotype g (by designation of Sabrosky
& Crosskey, 1969 : 40), NEw SoutH WALEs: Yantabulla (BMNH, London) [examined]. —
OLD.

Genus PSEUDOGONIA Brauer & Bergenstamm

Pseudogonia Brauer & Bergenstamm, 1889 : 100 (32). Type-species: Gonia cinevascens Rondani,
1859 [=Tachina rufifrons Wiedemann, 1830], by monotypy. (EUROPE).

Gaediogonia Townsend, 1927a: 71. Type-species: Gaediogonia jacobsoni Townsend, 1927
[=Tachina rufifrons Wiedemann, 1830], by original designation. (SUMATRA).

rufifrons Wiedemann, 1830 : 318 (Tachina). Holotype 9, CxHina (UZM, Copenhagen)
[examined].—A.C.T.; NEw GUINEA; SOLOMOoNS; widespread ORIENTAL & southern
PALAEARCTIC REGIONS; AFRICA.
cimevascens Rondani, 1859 : 34 (Gonia). Syntypes 3 4, 49 [unlabelled], Iraty (MZ, Florence).
jacobsont Townsend, 1927a : 71 (Gaediogonia). Holotype g, SUMATRA: Fort de Kock (ZM,
Amsterdam) [examined].
(For other synonyms see Crosskey, 1966a : 678)

Tribe ERYCIINI Robineau-Desvoidy
ERYCINAE Robineau-Desvoidy, 1830 : 142. Type-genus: Evycia Robineau-Desvoidy, 1830.

Genus APLOMYA Robineau-Desvoidy

Aplomya Robineau-Desvoidy, 1830 : 184. Type-species: Aplomya zonata Robineau-Desvoidy,
1830 [=Tachina confinis Fallén, 1820], by subsequent designation of Robineau-Desvoidy
(1863 : 459). (EUROPE).

Leiosia Wulp, 1893: 185. Type-species: Leiosia flavisquama Wulp, 1893, by monotypy.
(Java). Syn. n.

flavisquama Wulp, 1893 : 186 (Leiosia). Lectotype gf (by designation of Crosskey, 1969 : 104),
Java (ZM, Amsterdam) [examined]. Comb. n.-—Q.p; Java, Maraya, INDIA.

Undetermined sp. — S.A.

Genus APROTHECA Macquart

Aprotheca Macquart, 1851 : 148 (175). Type-species: Aprotheca vufipes Macquart, 1851, by
monotypy.

Parvabrachelia Townsend, 1916c¢ : 159. Type-species: Masicerarufipes Macquart, 1847 [= Myobia
tenuisetosa Macquart, 1847], by original designation. Syn. n.

vufipes Macquart, 1851 : 149 (176). Lectotype ¢ (by fixation of Townsend, 1932 : 49),
TasMANIA (MNHN, Paris) [examined].—Tasm. (Probably = tenwisetosa)

With the new assignment of Masicera vufipes Macquart, 1847, to Aprotheca the name
vufipes Macquart, 1851, becomes a junior secondary homonym. No replacement name is
proposed as future study will probably show that the types are conspecific, thus obviating
the need for a new name.

tenuisetosa Macquart, 1847 : 90 (74) (Myobia). Holotype g, Tasmania (BMNH, London)
[examined]. Comb. n.— Tas.
vufipes Macquart, 1847 : 86 (70) (Masiceva). Holotype g¢, Tasmania (BMNH, London)
[examined].

TACHINIDAE OF AUSTRALIA 155

Genus AUSTRONILEA Crosskey

Austronilea Crosskey, 1967b : 32. Type-species: Austronilea livida Crosskey, 1967, by origina
designation.

livida Crosskey, 1967b : 33. Holotype gj, AUSTRALIAN CAPITAL TERRITORY: Black Mt (ANIC,
Canberra) [examined]. — A.C.T.

Genus AUSTROPHRYNO Townsend

Austrophryno Townsend, 1916c : 160. Type-species: Tachina densa Walker, 1852 [=Evxorista
diversicoloy Macquart, 1847], by original designation.

Archimera Mesnil, 1954b : 371 (as subg. of Platymya Robineau-Desvoidy). Type-species:
Platymyia (Archimera) oncoperae Mesnil, 1954 [=Evxorista diversicoloy Macquart, 1847],
by monotypy.

diversicolor Macquart, 1847 : 83 (67) (Exorista). Holotype 3, Tasmania (BMNH, London)
[examined]. — A.C.T., N.S.W., Tasm.
densa Walker, 1852 : 288 (Tachina). Holotype 9, New SoutH Wares (BMNH, London)

[examined].

hebes Walker, 1852 : 289 (Tachina). Holotype g [not 9], Tasmania (BMNH, London)
[examined].

evythropus Walker, 1861c : 298 (Nemoraea). Holotype g, Tasmania (BMNH, London)
[examined].

oncoperae Mesnil, 1954b : 371 (Platymyia (Archimera)). Holotype g, Tasmania (CNC,
Ottawa, ex coll. Mesnil) [examined].

Genus BACTROMYIELLA Mesnil

Bactromyiella Mesnil, 1952 : 239-240. Type-species: Bactromyiella aureocincta Mesnil, 1952
[=Masicera ? ficta Walker, 1861], by original designation. (FIJ1).

ficta Walker, 1861b : 286 (Masiceva ?). Holotype 9 [head lost], InponEs1a: Moluccas, Batjan
(publ. as ‘Batchian’) (BMNH, London) [examined]. — Qtp; NEw GuINEA, Fij1, INDONESIA,
_ INDIA.
semivufa Malloch, 1930b : 351 (Stuymia). Holotype g, QUEENSLAND: Kuranda (ANIC,
Canberra) [examined].
auveocincta Mesnil, 1952 : 240 (Bactromyiella). Holotype g, Fiy1: Nadala (BMNH, London)
[examined].

Genus CHLOROGASTROPSIS Townsend

Chlorogastrvopsis Townsend, 1926c : 544. Type-species: Chlovogastey rvufipes Schiner, 1868
[=Tachina orga Walker, 1849], by original designation. (‘NEw ZEALAND’ in error).
Eipogonoides Curran, 1938: 195. Type-species: LEipogonoides vuficornis Curran, 1938

[=Tachina orga Walker, 1849], by original designation. Syn. n.

orga Walker, 1849 : 752 (Tachina). Lectotype g (by designation of Crosskey, 1966b : 108),
New SoutH WaLEs: Sydney (BMNH, London) [examined]. Comb. n.—N.S.W., OLD.
vufipes Schiner, 1868 : 323 (Chlovogaster). Lectotype g (by present designation), [AUSTRALIA:
publ. as ‘New Zealand’ in error] (NM, Vienna) [examined]. Syn. n.
Published as from New Zealand, but correct provenance given as ‘Neuholland’
(=Australia) by Brauer & Bergenstamm (1889 : 97).
vuficovnis Curran, 1938 : 196 (Eipogonoides). Holotype g, NEw SoutH Wares (BMNH,
London) [examined].

156 RE We CROSSE N

Genus ERYTHROCERA Robineau-Desvoidy

Erythrocera Robineau-Desvoidy, 1848 : 436. Type-species: Phryno nigripes Robineau-Desvoidy
1830, by subsequent designation of Robineau-Desvoidy (1863 : 600). (EUROPE).

facialis Mesnil, 1952 : 253. Holotype g, QuEENSLAND: Herberton (DEI, Eberswalde)
[examined]. — A.C.T., OQLp.

Genus METAPHRYNO Crosskey

Metaphryno Crosskey, 1967b : 28. Type-species: Metaphryno bella Crosskey, 1967, by original
designation.

bella Crosskey, 1967b : 29. Holotype ¢, NEw SoutH WaLEs: 3 mls NW of Rules Point
(ANIC, Canberra) [examined]. — N.S.W., Vict.

Genus PHOROCEROSTOMA Malloch

Phorocerosoma Malloch, 1929b : 327. Type-species: Phorocerosoma setiventris Malloch, 1929,
by original designation. [Junior homonym of Phorocerosoma Townsend, 1927.]
Phorocerostoma Malloch, 1930) : 326. [Replacement name for Phorocerosoma Malloch.]

setiventre Malloch, 1929b : 327 (Phorocerosoma setiventris). Holotype g, QUEENSLAND:
Macpherson Range, National Park (AM, Sydney) [examined]. — N.S.W., QLD.

Genus PSEUDALSOMYIA Mesnil

Pseudalsomyia Mesnil, 1968 : 178. Type-species: Pseudalsomyia piligena Mesnil, 1968, by
original designation. (PAKISTAN).

pilifacies Mesnil, 1968 : 180. Holotype g, NEw SoutuH WatEs: Lisarow (BMNH, London)
[examined]. — N.S.W.

Genus TERETROPHORA Macquart

Teretvophova Macquart, 1851 : 174 (201). Type-species: Tevetrophora fasciata Macquart, 1851,
by monotypy.

fasciata Macquart, 1851 : 175 (202). Holotype 9, ‘Tasmanta’ (MNHN, Paris) [examined]. —
N.S.W., Tasm. (?).
Undescribed sp. — N.S.W.

Unplaced species of Eryciini

crassiseta Baranov, 1938) : 409 (Bactromyia). Holotype 9, QUEENSLAND: Biloela (BMNH,
London) [examined]. — OLp.

quadrisetosa Curran, 1938 : 204 (Zenillia). Holotype 9, QUEENSLAND: Palm Is. (SPHTM,
Sydney) [examined]. — OLp.

varipes Macquart, 1846 : 291 (163) (Masiceva). Holotype g [labelled ‘Exorista varipes’],
Tasmania (MNHN, Paris) [examined]. — Tasm.

This species, referred to as Phryno varipes by Robineau-Desvoidy (1863 : 543), is
apparently near Austrophryno diversicolor (Macquart) but differs by having the parafacials
entirely bare and is left generically unassigned until it can be studied more fully.

Undescribed genera & spp. — N.S.W., S.A., Tas.

TACHINIDAE OF AUSTRALIA 157

Unplaced species of Tachinidae

armiceps Malloch, 1930b : 336 (Voriella). Holotype 9, WESTERN AUSTRALIA: Eradu (ANIC,
Canberra) [examined]. — W.A.

calliphon Walker, 1849 : 777 (Tachina). Holotype g [with puparium], ‘Picton’ (BMNH,
London) [examined].

Austen (1907 : 339) noted that the type bears a label ‘Picton’ and that it is presumably
therefore from ‘either New South Wales or Canada’. The type also bears (as Austen
did not note) a BMNH accession label reading ‘47 109’ (i.e. the 1ogth collection of insects
registered as received by BMNH in the year 1847). Examination of the register shows
that this collection consisted of many miscellaneous insects from ‘W. Australia’, and
suggests that calliphon must have an Australian provenance; pending other evidence
(from later-collected material of the same species which will help to pinpoint the locality)
the provenance of the holotype is accepted as Picton, Western Australia (though Picton,
New South Wales is an alternative possibility). Up to now the holotype has remained
unique; no specimens have been found that associate with it.

despicienda Walker, 1861c : 306 (Tachina). Holotype 9 [bad condition], NEw SoutH WALES
[?] (BMNH, London) [examined].

This species was published as from New South Wales and the holotype bears an old ink
label ‘NSW’. No Australian specimens have been seen that associate with the holotype,
which may not have had an Australian provenance. From the surviving characters of
the holotype it appears to belong near the genus Mauritiodoria Townsend from Mauritius
and might be the female of Mauritiodoria spinicosta (Thomson).

diversa Walker, 1852 : 262 (Ocyptera ?). Holotype 9, ‘TAsMANIA’ (lost).

This name remains a nomen dubium. The holotype is lost, and evidence that it originated
from Tasmania is inconclusive (there are doubts about several of the provenances cited
by Walker in the 1852 work).

hyalipennis Macquart, 1855 : 122 (102) (Phoroceva). Type(s) g, SourH AusTRALIA: Adelaide
(lost).-S.A. (Nomen dubium).

As the type-material is lost this name remains completely enigmatic; it is a junior

primary homonym of Phorocera hyalipennis Macquart, 1851, from Java.

inconspicua Malloch, 1930b : 336 (Voriella). Holotype 9, New SoutH Waters: Sydney
(SPHTM, Sydney) [examined]. — N.S.W.

lateralis Macquart, 1851 : 176 (203) (Degeeria). Holotype g, Tasmania [publ. as ‘Oceania’]
(MNHN, Paris) [examined].—Tasm. (Nomen dubium).

The holotype of this species is in appalling condition, being wholly coated with a brittle
deposit and completely concealed in mould. The name therefore remains enigmatic;
it is a junior primary homonym of Degeeria lateralis Macquart, 1848, from North America.

melas Bigot, 1889 : 256 (Exorista). Holotype 9, TAsMANIA (lost). (Nomen dubium).

The holotype of this species was not in Bigot’s collection when that collection came
to the BMNH and has not been seen since the time of description. Bigot added the word
‘Detrita’ after his Latin description, and the holotype was presumably therefore in very
bad condition when described. The name remains enigmatic.

mucrocornis Macquart, 1851 : 174 (201) (Phoroceva). Holotype 9, ‘Tasmania’ (MNHN,
Paris) [examined].— Tasm. (?). (? Blondeliini or Exoristini).

The holotype of this nominal species is in such bad condition that few features can be

made out. It appears, however, to be fairly certainly either a blondeliine or an exoristine).

SUMMARY OF NOMENCLATURAL CHANGES ESTABLISHED
IN THE CATALOGUE
The nomenclatural changes established in the foregoing catalogue are summarized
below in their appropriate categories. The order is alphabetical and in the tables
of synonyms the invalid junior names are cited first.

158 R. W. CROSSKEY

(a) New synonymy in genus-group names

Austrodexia Malloch, syn. n. of Senostoma Macquart.
Chlorogastrina Crosskey, syn. n. of Tasmaniomyia Townsend.
Dicephalomyia Malloch, syn. n. of Senometopia Macquart.
Eipogonoides Curran, syn. n. of Chlovogastvopsis Townsend.
Eocarcelia Townsend, syn. n. of Senometopia Macquart.
Eocarceliopsis Townsend, syn. n. of Senometopia Macquart.
Lasiocalyptey Malloch, syn. n. of Senostoma Macquart.
Lastocalyptrina Malloch, syn. n. of Senostoma Macquart.
Leiosia Wulp, syn n. of Aplomya Robineau-Desvoidy.
Pavabrachelia Townsend, syn. n. of Aprotheca Macquart.
Rhynchiodexia Bigot, syn. n. of Senostoma Macquart.

Ruya Paramonov, syn. n. of Rutilotyixa Townsend.
Schizactiana Curran, syn. n. of Cevomya Robineau-Desvoidy.
Schizoceromyia Townsend, syn. n. of Cervomya Robineau-Desvoidy.

(b) New synonymy in species-group names

Bessevioides sexualis Curran, syn. n. of Besserioides varicoloy (Curran).

Chlorogastey vufipes Schiner, syn. n. of Chlorogastropsis orga (Walker).

Exorista lata Macquart, syn. n. of Winthemia lateralis (Macquart).

Exorista marginata Macquart, syn. n. of Winthemia lateralis (Macquart).

Exorista vufomaculata Macquart, syn. n. of Phorocerosoma cilipes (Macquart).
Heterometopia vufipalpis Macquart, syn. n. of Hetervometopia argentea Macquart.
Linnaemyia nigripalpus Tryon, syn. n. of Linnaemya concavicornis (Macquart).
Machrochloria calliphorosoma Malloch, syn. n. of Machrochloria nitidiventris (Macquart).
Macrochloria caliphorosoma v. vufipes Malloch, syn n. of Macrochloria nitidiventris (Macquart).
Masiceva consanguinea Macquart, syn. n. of Exorista flaviceps Macquart.
Microtropeza fallax Hardy, syn. n. of Microtropesa violacescens Enderlein.
Nemorvaea brevisetosa Macquart, syn. n. of Winthemza lateralis (Macquart).
Ocypteva flavifrons Macquart, syn. n. of Cylindromyia bimacula (Walker).
Omalogaster limbinevris [sic] Macquart, syn. n. of Hetervometopia argentea Macquatt.
Omalogaster nitidus Macquart, syn. n. of Heterometopia argentea Macquart.
Phorocera acutangulata Macquart, syn. n. of Paropsivora graciliseta (Macquart).
Phoroceva maculata Macquart, syn. n. of Blepharella lateralis Macquart.

Phoroceva subpubescens Macquart, syn. n. of Polychaeta nigra Macquart.

Prosena albifrons Malloch, syn. n. of Prosena conica Guérin-Méneville.

Prosena indecisa Malloch, syn. n. of Pvosena macyopus Thomson.

Prosena parva Malloch, syn. n. of Prosena dorsalis Macquart.

Senostoma punctum Walker, syn. n. of Senostoma appendiculatum (Macquart).
Tachina australis Walker, syn. n. of Blepharipa fulviventris (Macquart).

Winthemia albiceps Malloch, syn. n. of Winthemia lateralis (Macquart).

Winthemia diversa Malloch, syn. n. of Winthemia neowinthemioides (Townsend).
Zosteromyia longicornis Hardy, syn. n. of Zosteromeigenia mima Townsend.

(c) New combinations*

[Note: The new combinations shown are only those that are considered taxonomically valid.
The list excludes combinations implied by new synonymy. ]

Alophova (Alophorella) chrysis (Malloch) comb. n.
Alophova (Alophorella) costalis (Malloch) comb. n.
* See also Appendix, p. 209.

TACHINIDAE OF AUSTRALIA

Alophora (Alophorella) discalis (Malloch) comb. n.
Alophora (Hyalomya) normalis (Curran) comb. n.
Alophora (Mormonomyia) basalis (Malloch) comb. n.
Alophova (Mormonomyia) hyalis (Malloch) comb. n.
Alophora (Mormonomyia) lativentris (Malloch) comb. n.
Alophora (Mormonomyia) lepidofera (Malloch) comb. n.
Alophora (Mormonomyia) nigrihirta (Malloch) comb. n.
Alophora (Mormonomyia) sensua (Curran) comb. n.
Alophora (subg. indet.) hippobosca (Paramonov) comb. n.
Alophora (subg. indet.) nigyisquama (Malloch) comb. n.
Anagonia anguliventris (Malloch) comb. n.
Anagonia grisea (Malloch) comb. n.

Anagonia lasiophthalma (Malloch) comb. n.
Anagomia lateralis (Macquart) comb. n.

Anagonia majoy (Malloch) comb. n.

Anagonia opaca (Malloch) comb. n.

Anagonia scutellata (Malloch) comb. n.

Apatemyia flavipes (Macquart) comb. n.

Apatemyia vufiventris (Macquart) comb. n.

Aplomya flavisquama (Wulp) comb. n.

A protheca tenuisetosa (Macquart) comb. n.
Argyrothelaiva melancholica (Mesnil) comb. n.
Blepharipa auripilis (Robineau-Desvoidy) comb. n.
Blepharipa coesiofasciata (Macquart) comb. n.
Blepharipa fulviventyis (Macquart) comb. n.

Ceracia aymata (Malloch) comb. n.

Ceracia fergusoni (Malloch) comb. n.

Ceromya fergusoni (Bezzi) comb. n.

Ceromya valida (Curran) comb. n.

Chaetophthalmus ruficeps (Macquart) comb. n.
Chlorogastropsis orga (Walker) comb. n.

Cuphocera pilifacies (Macquart) comb. n.

Eozenillia remota (Walker) comb. n.

Ethilla translucens (Macquart) comb. n.

Gerocyptera tristis (Bigot) comb. n.

Hillomyia polita (Malloch) comb. n.

Linnaemya concavicornis (Macquart) comb. n.
Macrochloria nitidiventris (Macquart) comb. n.
Medinodexia morgani (Hardy) comb. n.

Paropsivora australis (Macquart) comb. n.
Paropsivora graciliseta (Macquart) comb. n.
Paropsivora tessellata (Macquart) comb. n.

Peribaea argentifrons (Malloch) comb. n.

Peribaea baldwimi (Malloch) comb. n.

Peribaea orbata (Wiedemann) comb. n.

Peribaea plebeia (Malloch) comb. n.

Phorocerosoma cilipes (Macquart) comb. n.

Pilimyia lateralis (Macquart) comb. n.

Rutilotrixa diversa (Paramonov) comb. n.
Rutilotvrixa monstruosa (Paramonov) comb. n.
Rutilotvixa westralica (Paramonov) comb. n.
Senostoma apicale (Curran) comb. n.

Senostoma appendiculatum (Macquart) comb. n.
Senostoma atripes (Malloch) comb. n.

159

160 R. W. CROSSKEY

Senostoma basale (Curran) comb. n.

Senostoma brevipalpe (Macquart) comb. n.
Senostoma brevipalpe (Rondani) comb. n. (preocc. brevipalpe Macquart, see p. 117).
Senostoma commune (Malloch) comb. n.
Senostoma flavohivtum (Malloch) comb. n.
Senostoma hivticauda (Malloch) comb. n.
Senostoma hyria (Walker) comb. n.

Senostoma mixtum (Malloch) comb. n.
Senostoma modestum (Malloch) comb. n.
Senostoma nigrvihivtum (Malloch) comb. n.
Senostoma notatum (Walker) comb. n.
Senostoma pallidihivtum (Malloch) comb. n.
Senostoma punctipenne (Macquart) comb. n.
Senostoma rubricarinatum (Macquart) comb. n.
Senostoma setigerum (Malloch) comb. n.
Senostoma setiventve (Malloch) comb. n.
Senostoma taylori (Curran) comb. n.

Senostoma tessellatum (Macquart) comb. n.
Senostoma testaceicorne (Macquart) comb. n.
Senostoma untpunctum (Malloch) comb. n.
Sipholeskia cevtema (Curran) comb. n.
Tasmaniomyia tasmanensis (Macquart) comb. n.
Trigonospila braueri (Townsend) comb. n.
Trigonospila brevifacies (Hardy) comb. n.
Trigonospila fasciata (Hardy) comb. n.
Tritaxys scutellata (Macquart) comb. n.

(d) New names for junior homonyms

Hillomyia Crosskey nom. n., for Hillia Malloch (preoccupied by Hillia Grote).
macquarti Crosskey nom. n., for Masiceva auriceps Macquart, 1851 (preoccupied by Masicera
auriceps Macquart, 1843).

LECTOTYPE DESIGNATIONS

New lectotype designations are made below for twelve nominal species occurring
in Australia (ten described from Australia and two with an extra-Australian original
provenance). Each lectotype and available paralectotype has been appropriately
labelled.

Ceromasia sphenophori Villeneuve, 1911 : 81-82.

Described from one 9 and three $ syntypes sent to Villeneuve by de Meijere
for study. LECTOTYPE g, NEw Guinea: Papua [Laloki River area near Port
Moresby], vii-vili. 1909 (F. Muir) (in Zodlogisch Museum, Amsterdam).
Paralectotype gj, NEw GuINEA [probably same provenance as lectotype] (CNC,
Ottawa, ex Villeneuve-Mesnil coll.).

The lectotype bears labels in Villeneuve’s writing that read ‘Type’ and
‘Ceromasia sphenophori ¢ Villen’, and the paralectotype bears a Villeneuve label
reading ‘Ceromasia sphenophori type 3 Villen.’.

TACHINIDAE OF AUSTRALIA 161

The ZM, Amsterdam, collection contains two 2 specimens of C. sphenophori
from Ambon (=Amboyna, Moluccas) each with a pencilled label, one reading
‘Ambon’ and the other ‘Ambon vi vu 08’. It is known that Muir collected in
Amboyna and obtained sphenophori there in the year prior to that in which he
obtained the species at his Laloki river camp in Papua, and the year date 08
(=1908) fits with this. It is possible that one of the Amboyna females is an
original syntype, even though Villeneuve mentioned only New Guinea in the
description. Good evidence is lacking, however, and the Amboyna specimens
are considered not to have any type-status (neither bears a label by Villeneuve).

Chlorogaster rufipes Schiner, 1868 : 323.

Described from one 9 and two ¢g syntypes. LECTOTYPE 4g, Ausrratia (in
Naturhistorisches Museum, Vienna). Paralectotype 9, data as lectotype (also in
NM, Vienna). The third (g) syntype has not been seen. The lectotype bears a
label in purple ink reading ‘rufipes Type Br. Bgst.’, another label reading
‘N. Holland Alte Sammlung’ and a third label reading ‘rufipes det. B.B.’ (the
specific name in ink, remainder in print); the paralectotype has identical labels.

The provenance was published originally as ‘Neuseeland’ but this was corrected
to ‘New Holland’ by Brauer & Bergenstamm (1889 : 97).

Dexia aditha Walker, 1848 : 854.

The type-material of this nominal species was cited as ‘Holotype’ in the recent
revision of the Rutiliini (Crosskey, 1973). Whilst that paper was in press, however,
it was discovered that a second original specimen had been inadvertently
incorporated into the Australian National Insect Collection (having been loaned to
the late Dr Paramonov several years ago and not returned to British Museum
(Natural History)). It is now certain that the original material consisted of two
syntypes, and the specimen earlier referred to as ‘holotype’ is here newly
designated as lectotype.

LECTOTYPE ¢, AusTRALIA: Western Australia, Swan River (Richardson) (in
British Museum (Natural History), London). Paralectotype 3, same data as
lectotype (also in BMNH).

Lectotype and paralectotype bear identical labels reading as follows: circular
white label with ‘43 14’ on one side and “Swan River’ on the reverse side in slightly
faded black ink; pencilled label ‘Swan R. W. Australia. Dr. Richardson. 43.14.’
in Austen’s writing. (The figures ‘43 14’ refer to the 14th collection of insects
incorporated into the BMNH collection in 1843.)

Microtopeza [sic] violacescens Enderlein, 1937 : 441.

Described from three ¢ and two 92 syntypes. LECTOTYPE 4, AUSTRALIA:
Queensland, Herberton, 3700 ft, xii. rg10 (Dodd) (in Deutsches Entomologisches
c

162 R. W. CROSSKEY

Institut, Eberswalde). Paralectotypes: 2 3, 2 9, same data as lectotype (gf & 2
in DEI, Eberswalde, and ¢ & 2 in MNHU, Berlin).

Each type-specimen has a printed label ‘Herberton Dodd XII.1910 3700 Ft.’ and
Enderlein’s name label. The generic name is mis-spelt ‘Microtopeza’ on each name
label as well as in the original publication.

Palpostoma aldrichi Hardy, 1938 : 57 (= testacea sensu Aldrich).

Aldrich (1922) described a species of Palpostoma that he considered to be
P. testaceum Robineau-Desvoidy from a female and five male specimens from
Cairns in northern Queensland. Hardy (1938) considered that Aldrich’s species
could not be the true P. testacewm, as Robineau-Desvoidy would not have had the
same species. This supposition of Hardy is the merest conjecture, unsupported
by any real evidence (as Robineau-Desvoidy’s original material of testaceum is
lost and it is still not known, in the unsatisfactory taxonomic state of Palpostoma,
whether the same species may occur in north Queensland as are found in New
South Wales—the most probable provenance of Robineau-Desvoidy’s material);
nevertheless, Hardy’s name P. aldrichi that he published for Aldrich’s supposedly
misidentified testaceum is available in nomenclature. The name is not accompanied
by any description, but the reference is given to Aldrich’s (1922) description under
the name ¢estacea [sic] and the name aldrichi is therefore available under Article 13
(a) (ii) of the ICZN.

The type-material of aldrichi is comprised of the six specimens cited by Aldrich.
Aldrich stated that three of these (¢) had been returned to the collector (Illingworth),
and the others (two g, one 9) retained for the USNM collection. The specimens
returned to Australia have not been located (though possibly still present in a
collection in Brisbane) but the others are in Washington, and a lectotype is here
designated from them.

LECTOTYPE g, AusTRALIA: Queensland, Cairns, 1919 (J. F. Illingworth) (in
United States National Museum, Washington D.C.). Paralectotype ¢, same data
as lectotype (label lacking collector’s name) (USNM); paralectotype 2, Queensland,
Babinda, 1919 (J. F. Illingworth) (USNM).

The lectotype is labelled “Cairns N. Q. 19’ and ‘J. F. Illingworth Coll. Ex.
Window’ and bears Aldrich’s identification label as ‘testacea’. There are
discrepancies between the labelling of the paralectotypes and Aldrich’s published
data, but it seems probable that Aldrich was citing data only from the specimen
that he retained and labelled (i.e. the lectotype) and that the two specimens here
treated as paralectotypes that he also retained (a gj and a 9) are part of the type-series
in spite of the discrepancies; the female is labelled as from Babinda (not Cairns)
and the male is labelled as collected by A. P. Dodd (not J. F. Illingworth).

Senostoma ? punctum Walker, 1858 : 205.

Described from ‘Australia and New South Wales’, from which statement it is
clear that there were at least two original syntypes. Two specimens are in

TACHINIDAE OF AUSTRALIA 163

BMNH collection, one from New South Wales and the other without locality label;
these are considered to be both original syntypes as both came from Saunders’
collection (they are males, whereas Walker stated ‘Female’, but errors of sexing
are frequent in Walker’s work). LECTOTYPE 4, Austratia: New South Wales
(in British Museum (Natural History), London). Paralectotype 3, AUSTRALIA
presumed (also in BMNH, London).

Sisyropa cinerea Brauer & Bergenstamm, 1891 : 346 (42).

Described from both sexes but without statement of the number of specimens.
Existing material consists of one 3 and one 2 syntype (mis-associated with each
other), LECTOTYPE 3, Australia: Queensland, Rockhampton, 1868 (Thorey)
(in Naturhistorisches Museum, Vienna). (Only the Rockhampton locality is
mentioned in the original description.)

The lectotype bears a label reading ‘Thorey 1868 Rock-hampton’ (the word
Thorey and figures 186 in print, remainder in black ink), a label ‘cinerea det. B. B’
(handwritten in black ink except for the letters ‘det.’) and a label in purple ink
reading ‘Blepharipoda Sisyropa’. The paralectotype is similarly labelled except
that the words ‘Cap York’ are given for locality on the first label and there is no
label in purple ink.

Lectotype and paralectotype are mis-associated, the former being a Carcelia
and the latter a winthemiine (Nemorilla sp.). Mesnil (19500: 9) referred to ‘le
Carcelia (Eucarcelia) cinerea B. B. dont nous avons vu le type’, but this statement
does not restrict the name to a single recognizable specimen and therefore does not
constitute a valid lectotype fixation. Present designation of the g specimen as
lectotype maintains the sense of the name used by Mesnil. Here it should be
noted that Crosskey (19660 : 109) cited his assignment of cinerea to Carcelia as a
new combination; this was due to an oversight, Mesnil’s (19500) earlier assignment
to Carcelia in a work on African Carceliini having been overlooked.

Tachina bura Walker, 1849 : 760.

Described from two specimens, one presented to BMNH by Rev. T. Ewing
from Van Diemen’s Land, and the other from New Holland and in the Rev. J.
Wenham’s collection. Only the specimen from Tasmania has been found, which
is here designated as lectotype. LECTOTYPE gd, AvustTraia: Tasmania (in
British Museum (Natural History), London).

The lectotype bears a circular white accession label with the ink figures ‘46 81’
and a pencilled label in Austen’s writing that reads ‘Tasmania. Rev. J. Ewing.

46. 81’.

Tachina ruralis Fallén, 1810 : 265.

Described from an unstated number of specimens of both sexes from ‘Esperéd i
Skane’. The Fallén collection at Stockholm contains seven specimens standing

164 R. W. CROSSKEY

under the name ruralis, none of them with data labels; one is 3 and six are 9.
In the absence of contrary evidence all seven specimens are accepted as syntypes.
The g syntype is the only specimen that belongs to the well-known and nearly
cosmopolitan species that has long been known as Vora ruralis (Fallén) and this
specimen is therefore designated as lectotype. LECTOTYPE 3, SwEepen: Skane,
Esperéd (in Naturhistoriska Riksmuseum, Stockholm). Paralectotypes 7 2, data
presumed as lectotype (also in NR, Stockholm).

The lectotype bears a very old label reading (in faded ink) “Tachina ruralis 3
Fallén’, and one of the @ paralectotypes bears an identical label. Other
paralectotypes are unlabelled.

All six 9 paralectotypes are mis-associated with the g lectotype. They belong
to a species of Exorista, most of them probably being Exorista rustica (Fallén,
1810). This species was described immediately before ruralis in Fallén (1810 : 264),
and Fallén noted in the description of ruralis its similarity to rustica. There
may at some time have been a confusion of the specimens of rustica and ruralis,
but the present designation of the g syntype as lectotype maintains the
longstanding usage of the name ruralis for a species of Voria Robineau-Desvoidy
and not of Exorista Meigen.

Zosteromyia brevifacies Hardy, 1934 : 36.

Described from one 2 syntype from Tooloom (N.S.W.) and a 9 and two ¢
syntypes from Mt. Wellington (Tasmania). The Tooloom specimen is designated
as lectotype; the other syntypes have not been located. LECTOTYPE 9Q,
AUSTRALIA: New South Wales, Tooloom, 29.i.1926 (in Queensland Museum,
Brisbane: registered No. T.7127).

The lectotype bears an ink label reading ‘Tooloom NSW 29.1.26’ and a name
label in Hardy’s writing reading ‘Zosteromyia brevifacies Hardy PARATYPE’.
(As no holotype was designed in the original publication the specimen is an original
syntype, not a paratype.)

Zosteromyia longicornis Hardy, 1934 : 36.

Described from three ¢ syntypes from Brisbane and Mt _ Glorious.
LECTOTYPE g, AUSTRALIA: Queensland, Mt Glorious, 25.iv.1930 (in University
of Queensland, Brisbane). Paralectotype 3, same data as lectotype (in BMNH,
London). The third syntype has not been located.

The lectotype bears an ink label reading ‘Mt Glorious 25.4.30’ and a name label
in Hardy’s writing reading ‘Zosteromyia longicornis Hardy PARATYPE’. (As
no holotype was designated in the original publication the specimen is an original
syntype, not a paratype.)

Zosteromyia morgani Hardy, 1934 : 37.

Described from four ¢ specimens (syntypes) without a designated holotype.
LECTOTYPE 4, AustRALiA: New South Wales, Biniguy [publ. as ‘Binniguy’],

TACHINIDAE OF AUSTRALIA 165

22.11.1930 (W. L. Morgan) (in New South Wales Department of Agriculture,
Rydalmere). Paralectotype: 1 3, same data as lectotype (in NSWDA, Rydalmere) ;
I g, New South Wales, Narara [publ. as ‘Worara’], I1.xi.193I [publ. as ‘30’]
(W. L. Morgan) (NSWDA, Rydalmere).

The lectotype bears an ink label reading ‘Bred from Aulacophora hilaris adult
coll. Binniguy 22.2.30. W. L. Morgan’ and the condition is fair except for some
collapse of the eyes and scutum and loss of apical half of left wing. One
paralectotype is labelled exactly as the lectotype and has its associated puparium
(the abdomen is gummed separately to the card mount); the other paralectotype
is labelled in ink ‘Bred from Aulacophora hilaris adult coll. Narara 11.11.31
W. L. Morgan’.

The lectotype and paralectotypes each bear a name label in Hardy’s writing
that reads ‘Zosteromyia morgani Hardy PARATYPE’ and a printed label
‘Department of Agriculture, Sydney, N.S.W., Australia’.

SUMMARY OF AUSTRALIAN NOMINAL SPECIES FOR WHICH TYPES
ARE LOST OR MISSING

The foregoing catalogue contains the names of 487 nominal species-group taxa
described from Australia (up to 1973). Primary types are known to exist for
458 of these taxa, but are lost or have not been located for the remaining 29 taxa.
The lost or missing types fall into two categories that are differentiated in the lists
of missing types that follow. The first category is that of types that can
confidently be asserted to be lost: in these cases (all of them nominal species
described by nineteenth-century authors) the types have never been found by later
workers and no specimens that could be the types have been found during personal
searches of likely depositories carried out for the preparation of this catalogue.
The second category is that of types which have not been found during the
preparation of this work but which may still exist: in these cases (mainly nominal
species described by twentieth-century authors) it is probable that types still exist
and will eventually be found in some small collection or unexpected place (those
of the two species described by Rondani, for example, may well be in a small
Italian museum but appear certainly not to be in Florence, Naples, or Genoa).
The distinction here made between ‘lost’ and ‘missing’ types is emphasized to
assist future revisionary work: it is considered that later workers may safely
assume that the types listed as ‘lost’ are truly lost or destroyed and will never be
found; on the other hand, searches will need to be made for the types listed as
‘missing’ whenever the relevant groups are studied in detail. The lists are
alphabetical under the original binomina.

(a) Nominal species of which the types are lost

Carcelia tasmanica Robineau-Desvoidy
Dexia hyria Walker
Exorista melas Bigot

166 R. W. CROSSKEY

Micropalpus bicolor Macquart.

Musca sinuata Donovan

Ocyptera ? diversa Walker

Omalogaster nitidus Macquart
Palpostoma testacea Robineau-Desvoidy
Phorocera hyalipennis Macquart (1855)
Rutilia australasia Gray

Rutilia fuluvpes Guérin-Méneville
Rutilia vidua Guérin-Méneville
Verreauxia auripilis Robineau-Desvoidy

(b) Nominal species of which types are missing

Calopygidia castanea Hardy
Cuphocera pilosa Malloch
Dexia brevipalpis Rondani
Euthera skuset Bezzi

Exorista trichopareia Schiner
Linnaemyia nigripalpus Tryon
Miucrotropesa skusei Bergroth
Miucrotropeza fallax Hardy
Prosena albifrons Malloch
Prosena indecisa Malloch
Prosena varia Curran
Rhinomyobia australis Brauer & Bergenstamm
Rutilia spinolae Rondani
Schizotachina fergusoni Bezzi
Zosteromyia fasciata Hardy
Zosteromyia minor Hardy

PART III-A HOST CATALOGUE FOR THE AUSTRALIAN TACHINIDAE

INTRODUCTION

The hosts of very nearly all true Tachinidae (from which I exclude the
Rhinophoridae, a group sometimes treated as tachinids) are other insects, but
centipede hosts are known. As a rule the larval or pupal stages of the hosts are
parasitized, especially the caterpillars of Lepidoptera and Hymenoptera Symphyta
and the soil- or wood-inhabiting grubs of Coleoptera, but when hemimetabolous
insects are the hosts it is usually the adult stage that is attacked; a few forms
parasitize adult beetles. The host-relations of the Australian Tachinidae conform
in their essentials with the general picture of tachinid parasitism, and there are
no insect orders providing hosts in Australia that do not also provide hosts in
other zoogeographical regions. In Australia eight insect orders are so far known

TACHINIDAE OF AUSTRALIA 167

positively to provide hosts, and these same orders provide the hosts for the
overwhelming majority of world forms: the Lepidoptera and Coleoptera, as elsewhere,
provide the greatest number of different host species, and the Orthoptera,
Hemiptera-Heteroptera and Hymenoptera are regularly parasitized by particular
groups of Australian tachinids; a few species of stick-insects (Phasmatodea) and
mantids (Mantodea) also provide hosts. The rather well developed fauna of
Embioptera in Australia is not known to have tachinid parasites, but may be found
to do so (as the tachinid genus Rossimyiops Mesnil is parasitic on an embiopteran
in South Africa). There is one remarkable record of an Australian tachinid
parasitizing adult Tabanidae (Diptera) (Spratt & Wolf, 1972).

Tachinids are presumed to play an important role in the natural regulation
of the numbers of their hosts, but this is difficult to quantify. In Australia many
of the economically important insect pests are attacked by tachinid parasites,
and some parasite species are regularly reared in numbers from their host pest
species. Particular tachinid groups may be confined to particular host groups
(e.g. the Phasiinae only attack Hemiptera and the Acemyini only attack
Orthoptera) but true host-specificity in the sense of a single species of parasite
confined to a single host-species is apparently rare in the Australian fauna (and
the apparent instances where host-specificity occurs are probably mainly due to
insufficient knowledge). Certainly several of the main injurious pests are attacked
by several species of tachinid, and many of the tachinids attacking these pests
also have other hosts (as is evident from the accompanying parasite-host and
host-parasite lists).

The range of economically important Australian insect pests that are attacked
by Tachinidae is very diverse, and includes pests of agricultural crops (sugar-cane,
cotton, maize, cucurbits) and many serious defoliators of forest timbers. Some
of the most important pests, with their parasite-groups, are: the cotton bollworm
(Heliothis armigera), the army-worm (Persectania, Pseudaletia, Spodoptera spp.)
and cutworm (Agvotis) pests of cotton and other crops that are attacked by many
species of Tachininae and Goniinae; the sugar-cane borer weevil (Rhabdoscelus
obscurus) attacked by Lixophaga sphenophori; the white-grub larvae of melolonthine
beetles, especially the sugar-cane white-grub (Dermolepida albohirta) of the
Queensland canefields, attacked by species of Palpostoma and of Rutiliini; the
introduced scarabaeid beetle Heteronychus arator, a pest of maize in New South
Wales, that is attacked by species of Palpostoma; the chrysomelid beetle
Aulacophora hilaris that destroys the foliage and flowers of cucurbitaceous crops,
attacked by a species of Blondeliini; the cotton-stainer bug (Dysdercus sidae) of
Queensland, attacked by species of Phasiinae; the pergid sawfly defoliators of
Eucalyptus, attacked by several species of the Anagonia-Froggattimyia complex
(Blondeliini); the chrysomelid beetle Pavopsis atomaria, a pest of Eucalyptus,
attacked by several species of Blondeliini; and the stick-insect Didymuria violescens,
a serious defoliator of Eucalyptus in New South Wales, that is attacked by an
undescribed tachinid of very uncertain systematic position.

Despite the diversity of economically important insect pests, and the extent
of work undertaken on them by Australian departments of agriculture and

168 R. W. CROSSKE Y

forestry, there has not up to now been any published host-list for the Australian
Tachinidae. The only published host records available have been scattered in
original tachinid descriptions or cited haphazardly in departmental reports or in
accounts of particular pests, and these were the main sources for the relatively few
entries in W. R. Thompson’s A Catalogue of the Parasites and Predators of Insect
Pests concerned with Australian Tachinidae. Many of the earlier records existing
in these various publications cannot be relied upon, either because of changes
in the nomenclature of the hosts and parasites or because of misidentification,
especially of the tachinids, and for some time an up-to-date host catalogue for the
Australian Tachinidae has been needed that is based so far as possible on reliably
named hosts and parasites and on the latest information available.

The host catalogue here presented may not be exhaustive, as there are probably
some tachinid specimens scattered in Australian collections that were reared from
known hosts but have not been available during the present study. Nevertheless
the lists of hosts and parasites are sufficiently comprehensive to form useful basic
lists that can be gradually augmented as more evidence on the host-relations is
acquired.

A major difficulty in compiling dependable host-parasite lists is the unreliability
of the identifications. As a rule, material of the hosts is not kept in collections
with the reared Tachinidae so that confirmation of identity of both host and
parasite is difficult or impossible. In general, however, it is likely that the hosts
will have been correctly identified, since they are commonly well known pests and
often are conspicuous Lepidoptera whose specific identities are not in doubt (even
if the lepidopterists are in dispute about the generic placements). On the other
hand identities of tachinid parasites are likely to be wrong unless they have
been recently checked by a specialist on the group (and in some difficult groups
of tachinids even this is no gaurantee for every specimen). In preparing the
accompanying parasite-host and host-parasite lists it has been assumed that the
hosts have been correctly identified, but the tachinids have only been recorded
when: (1) they have been personally identified, (2) when the host record is from the
original type-material of the tachinid parasite, or (3) when published records,
other than the original descriptions, are undoubtedly based on correctly identified
Tachinidae. The last circumstance is relatively infrequent, and most host records
in the literature have been discounted because the identities cited for the tachinid
parasites are either wrong or suspect (for example, most of G. H. Hardy’s
identifications of Australian Tachinidae were based on guesswork from the
literature and in consequence were often in error: hence his published host records
have usually been discounted).

The information for the host catalogue derives largely from specimens in the
collection of the British Museum (Natural History). Many of these specimens
have been received from time to time as duplicate specimens submitted to the
Commonwealth Institute of Entomology for identification, usually by Australian
state departments of agriculture and forestry; for this reason the BMNH collection.
is more comprehensive than any other in Australian Tachinidae reared from known
hosts, and the host catalogue is almost as completely comprehensive as it is possible

TACHINIDAE OF AUSTRALIA 169

to make it at the present time (though, as aforesaid, a search of collections in
Australia will yield up a few additional records that have not been known to me
while preparing the present work).

A SYNOPSIS OF THE HOST-RELATIONS OF AUSTRALIAN TACHINIDAE

Hosts are known for almost a quarter of the described Australian tachinid
fauna, but as the described fauna probably does not represent more than about
a quarter or fifth of the actual number of species in Australia it is evident that
knowledge of the hosts is only very fragmentary at present. The following
comments summarize the host-relations for the different host orders and parasite
groups, so far as they can be generalized from what is already known.

Lepidoptera. This order provides the hosts for the great bulk of forms in the
Tachininae and Goniinae, but is not parasitized by any Phasiinae or Proseninae.
Both butterflies and moths are attacked, and 27 families are so far known to
provide tachinid hosts in Australia. Some lepidopterous species, especially in
the Noctuidae, are attacked by several species of Tachinidae, at least nine species
attacking the army-worm Pseudaletia unipuncta.

Coleoptera. This order is next in importance to the Lepidoptera in the number
of host members it provides, though up to now only four families are known to
be involved as hosts of Australian Tachinidae. Members of the order are attacked
by the Proseninae, Palpostomatini, several genera of Blondeliini and apparently
by Apatemyia (probably Leskiini) and Pseudalsomyia (Eryciini). The Proseninae
and Palpostomatini are confined to beetle hosts in the larval and adult stages
respectively and mainly attack Scarabaeidae.

Hemiptera. MHeteropterous land bugs are hosts of the Phasiinae only, and in
Australia this subfamily (on the limited evidence so far) is confined to hemipterous
hosts. Members of the Coreidae, Lygaeidae and Pyrrhocoridae provide the hosts
so far discovered, but the Australian Pentatomidae are almost certainly parasitized
also (as the genus Pentatomophaga has pentatomid hosts in Java and New
Guinea).

Orthoptera. Acridoidea of the families Acrididae and Eumastacidae are the
hosts of the Acemyini, and this tribe is confined to acridoid hosts. The most
polyphagous species of tachinid known in the Australian fauna, though its host
species are all acridids and eumastacids, is Ceracia fergusoni, which has been reared
from 28 host species (22 undescribed and the others named). Blackith (1967) has
discussed this species under the name M yothyria fergusoni, and this work is the only
paper of any note that has yet appeared on the biology of any Australian
Tachinidae. Hosts are not yet known in Australia for Phorocerosoma (tribe
Ethillini) or the Ormiini, but it is likely that these tachinids will be found to have
orthopterous hosts: Phorocerosoma is a parasite of Acridoidea in Japan and in
Africa, and the Ormiini are parasites of nocturnally active Tettigoniidae s.l.
wherever the hosts are known (New World, southern Europe, Fiji).

170 Re Wi. CROSSE

Hymenoptera. In Australia the Hymenoptera are parasitized only by a few
members of the subfamily Goniinae. The Anacamptomylini are parasites in the
nests of certain Vespoidea, and several species of the Froggattimyia-Zenargomyia
complex (Blondeliini) attack the larvae of certain sawflies (Pergidae and Argidae).

Phasmatodea. Although 130 species of Phasmatidae are known in Australia
only two of these are known to be hosts of Tachinidae. These two species are
attacked by an undescribed species of Tachinidae that represents an undescribed
genus of doubtful affinity (but possibly belonging near the Acemyini). This
tachinid is remarkable for its very strong downcurved hook-like ovipositor. The
hosts of the genus Mycteromyrella (? Ethillini) are not known in Australia, but
in the Solomon Islands species of Mycteromyiella attack stick-insects of the genera
Ophicrania Kaup and Megacrania Kaup; it seems likely that the Australian species
of Mycteromyiella might similarly attack phasmatids.

Mantodea. Only one of the 118 species of Mantodea in Australia has so far
been recorded as host of a tachinid, viz. Pseudomantis albofimbriata which has
been found parasitized by Exorista coras. Ordinarily the Lepidoptera are hosts
of exoristine tachinids, but the record of EF. covas from a mantid and of an
unidentified Exorista species from a mantid in Guadalcanal (latter obtained by
R. W. Paine) suggest that parasitism of mantids by exoristine tachinids is an
occasional phenomenon in a group that habitually parasitizes lepidopterous
caterpillars. In other regions the Mantodea are parasitized by other tachinid
tribes that are probably close relatives of the Exoristini, such as the Masiphyini
in the Neotropical Region and certain Ethillini in the Ethiopian Region.

Diptera. The only record of Diptera as hosts of Tachinidae in Australia is that
of Spratt & Wolf (1972).

The subfamilies and tribes of Australian Tachinidae and their host groups

The following synopsis is given to show at a glance the various host groups for
the subfamilies and tribes of Tachinidae known to be represented in Australia.
The subfamilies and tribes are listed in the systematic order adopted in the
taxonomic catalogue (Part II). For some tribes there are no Australian host
records yet available, in which case the host information given is derived from
extra-Australian records and annotated as appropriate.

Subfamily Tribe Host-group
PHASIINAE Trichopodini Hemiptera-Heteroptera
Phasiini Hemiptera-Heteroptera
Cylindromyiini Hemiptera-Heteroptera [apparently no
Australian records to date}
Leucostomatini Hemiptera-Heteroptera
Eutherini Hemiptera-Heteroptera Pentatomidae [no
Australian records to date]
PROSENINAE Prosenini Coleoptera (larvae)

(DEXIINAE) Rutiliini Coleoptera (larvae)

TACHININAE
(MACQUARTIINAE)

GONIINAE

TACHINIDAE OF AUSTRALIA I7I

Palpostomatini*
Myiotrixini*
Ormiuini*

Glaurocarini*

Campylochetini
Voriini
Thelairini

Minthoini
Nemoraeini
Leskiini
Ernestiini
Parerigonini*

Linnaemyini
Tachinini
Acemyini
Neaerini
Siphonini
Blondeliini

Exoristini

Ethillini

Winthemiini
Carceliini
Anacamptomyiini
Sturmiini

Goniini
Erycuini

Coleoptera (adults)

[Unknown]

Orthoptera Tettigoniidae s.]. {no Australian
records to date]

Orthoptera Tettigoniidae of Glaurocara,
Lepidoptera of Doddiana [no Australian
records to date]

Lepidoptera (larvae)

Lepidoptera (larvae)

Lepidoptera (larvae) [no Australian records
to date]

Lepidoptera (larvae)

Lepidoptera (larvae)

Lepidoptera (larvae)

Lepidoptera (larvae)

[Apparently unknown, no Australian
records to date]

Lepidoptera (larvae)

Lepidoptera (larvae)

Orthoptera Acridoidea

Lepidoptera (larvae)

Lepidoptera (larvae)

Coleoptera (larvae and adults), Hymenoptera
Symphyta (larvae), Lepidoptera (larvae)
Lepidoptera (larvae), Hymenoptera Symphyta
(larvae) [no Australian records to date],

Mantodea [very rarely]

Lepidoptera (larvae), Orthoptera Acridoidea
for Phorocerosoma {no Australian records
to date], Phasmatodea for Mycteromyiella
{no Australian records to date, genus
possibly not true ethilline]

Lepidoptera (larvae)

Lepidoptera (larvae)

Hymenoptera Vespoidea (larvae)

Lepidoptera (larvae), Hymenoptera Symphyta
(larvae) [rarely, no Australian records to
date]

Lepidoptera (larvae)

Lepidoptera (larvae), Coleoptera
Cerambycidae (larvae) for Pseudalsomyia,
Hymenoptera Symphyta (larvae) [rarely,
no Australian records to date], Diptera
Tabanidae (adults) for Bactromyiella

* The affinities of the tribes so marked are very obscure. Their placement in Tachininae is an interim
measure until the relationships can be more clearly determined. Sabrosky & Arnaud (1966), following
Townsend, place the Palpostomatini in Phasiinae and the Glaurocarini and Ormiini in Proseninae.

PARASITE-HOST LIST

The tachinid parasites cited in the list are arranged in alphabetical order of their
tribes, and alphabetically by genus and species within each tribe; the names used

172 K. W. CROSSKEY

are those considered valid in the taxonomic catalogue (Part II). The names of
hosts are those considered currently valid and are arranged alphabetically within
each host family; when two or more families are represented in the host list
pertaining to any parasite then each begins on a separate line. The order and
family of the host(s) are shown in parentheses after the host name(s), and the
following abbreviations are used for the host orders: COL., Coleoptera; HEM.,
Hemiptera; HYM., Hymenoptera; LEP., Lepidoptera; MANT., Mantodea; ORTH.,
Orthoptera; PHAS., Phasmatodea. Subgeneric names are omitted for both
tachinids and hosts. Authors’ names are omitted for both parasite and host
species: those of the Tachinidae can be found in the taxonomic catalogue
(Part II) and those for the hosts are given in the ‘host-parasite list’ (beginning
on p. 178).

The great majority of host records are derived from data on tachinid specimens
in the British Museum (Natural History) collection or from host data published
with the original tachinid descriptions, or from both, and these are the sources
of information unless a host name is annotated by an entry in square brackets.
Annotations in square brackets refer to host records that are accepted as correct,
either on the basis of a published record that is unexceptionable or on the basis
of tachinid specimens from known hosts examined and identified by me and
housed in an Australian departmental collection. In citing such collections the
following abbreviations are used: NSWDA, New South Wales Department of
Agriculture, Rydalmere; QDPI, Queensland Department of Primary Industries,
Brisbane; WARI, Waite Agricultural Research Institute, South Australia. These
abbreviations are only used when the collections to which they refer contain
species from known hosts that are not duplicated by specimens in the BMNH
collection; when specimens of any tachinid parasite from the same host are housed
both in the BMNH collection and in NSWDA, QDPI or WARI then no annotation
is given.

Whenever such information has been available the hosts are listed for
undescribed or undeterminable species as well as those for which specific identities
are known. Similarly, hosts are indicated for known tachinid parasites in instances
where the host identities are not fully known.

Tachinid Parasites Hosts
ACEMYINI
Ceracia fergusont Azelota diversipes, Chortoicetes terminifera, Macrotona
australis, Urnisa rugosa (ORTH., Acrididae) [Blackith,
1967]

Keyacris interpres, Keyacris marcida, Moraba amicult,
Moraba keyi, Moraba misilliformis, Moraba viatica,
and 22 undescribed spp. (ORTH., Eumastacidae)
[Blackith, 1967]
Ceracia spp. Coryphistes oe Gastrimargus musicus (ORTH.,
Acrididae

TACHINIDAE OF AUSTRALIA 173

ANACAMPTOMYIINI

Anacamptomyia nigriventris Polistes tasmaniensis, Polistes sp. (HYM., Vespidae)

Euvespwora decipiens
Koralliomyia sp.
Koralliomyia sp.

BLONDELIINI

Anagonia anguliventris
Anagonia lasiophthalma
Anagonia lateralis

Anagonia scutellata
Compsilura concinnata

Froggattimyia hirta

Froggattimyia nicholsoni
Froggattimyia tillyardi
Froggattimyia wentworthi
Lecanipa sp. (2)
Lixophaga sphenophori
Medinodexia morgani
Monoleptophaga caldwelli
Paropsivora australis
Paropsivora grisea
Paropsivora sp.
Trigonospila brevifacies

Zenargomyia mooret

CAMPYLOCHETINI

Elpe sp.

CARCELIINI

Carcelia cosmophilae

Carcelia murina
Carcelia noctuae

Carcelia sp.
Carcelia sp.
Carcelia sp.
Carcelimyia dispar

Polistes sp. (HYM., Vespidae)
Ropalidia marginata jucunda (HYM., Vespidae)
Polistes sp. (HYM., Vespidae)

Paropsis atomaria (COL., Chysomelidae)

Gonipterus scutellatus (COL., Curculionidae)

Bryachus squamicollis (COL., Curculionidae). Unidentified
chrysomelid larva (COL.)

Unidentified chrysomelid (COL.)

Anomis xanthindyma (QDPI), Brithys crini, Spodoptera
sp. [QDPI] (LEP., Noctuidae)

Doratifera vulnervans [NSWDA] (LEP., Limacodidae)

Archips austvalana [NSWDA] (LEP., Tortricidae)

Numerous other LEP. hosts in extra-Australian regions

Lophyrotoma sp. Perga glabra, Pterygophorus analis
(Malloch, 1934] (HYM., Pergidae)

Perga dorsalis (HYM., Pergidae)

Paropsis atomaria (COL., Chrysomelidae)

Unidentified pergid (HYM.)

Pterolocerva sp. (LEP., Anthelidae)

Rhabdoscelus obscurus (COL., Curculionidae)

Aulacophora hilavis (COL., Chrysomelidae)

Monolepta australis (COL., Chrysomelidae)

Chrysophtharta bimaculata (COL., Chrysomelidae)

Paropsis atomaria (COL., Chrysomelidae)

Paropsis sp.

Heliocausta hemiteles [NSWDA] (LEP., Oecophoridae)

Phthorimaea operculella (LEP., Gelechiidae)

Zenarge turnert (HYM., Argidae)

Ocinara lewinae Lewin [NSWDA] (LEP., Bombycidae)
Scoliacma bicolora (LEP., Arctiidae)

Acantholeucania loreyi, Achaea janata [QDPI], Anomis
evosa [ODPI], Anomis flava, Plusia argentifera,
Pseudaletia unipuncta (LEP., Noctuidae)

Ialmenus evagovas [QDPI] (LEP., Lycaenidae)

Graphium macleayanus (LEP., Papilionidae)

Anthela varia [NSWDA], Anthela sp. (LEP., Anthelidae)

Anomis flava, Anticarsia ivvovata, Euxoa vadians [QDPI]
Heliothis avmigera, Heliothis punctigera [QDPI],
Pseudaletia unipuncta [QDPI] (LEP., Noctuidae)

Orgyia anartoides [NSWDA] (LEP., Lymantriidae)

Tisiphone sp. (LEP., Nymphalidae)

Delias aganippe [NSWDA] (LEP., Pieridae)

Ochrogastey contravia, Ochrogastey sp. (LEP., Notodontidae)

Panacela lewinae [NSWDA] (LEP., Eupterotidae)

174

ERNESTIINI

Chlorotachina sp. n.

ERYCIINI

Aplomya sp. ? flavisquama
Aplomya sp.

Bactromyiella ficta
Bactromyiella ? ficta

Chlovogastropsis orga
Pseudalsomyia pilifacies
Teretrophora fasciata

Tevetvophora sp.
Undetermined genus (near
Chlovogastropsis) 1
Undetermined genus (near
Chlorogastropsts) 2
Undetermined genus
Undetermined genus

ETHILLINI

Ethilla translucens

Ethilla sp.

EXORISTINI

Austrophorocera sp.
Eozenillia remota

Exorista covas
Exorista curriet

Exorista flaviceps

Exorista mungomeryt

Exorista psychidivora
Exorista sorvbillans

Exorista spp.

RK. W. CROSSKEY

Hesperilla sp. (LEP., Hesperiidae)

Evysichton lineata lineata [QDPI] (LEP., Lycaenidae)

Nacaduba biocellata biocellata (LEP., Lycaenidae)

Nacoleia octasema, unidentified sp. (LEP., Pyralidae)

Dasybasis hebes, Dasybasis oculata (DIPT., Tabanidae)
[Spratt & Wolf, 1972]

Metura elongata (LEP., Psychidae)

Unidentified cerambycid (COL.)

Avachnographa micrastrella, Heliocausta hemiteles [NSWDA\],
Philobota facialis (LEP., Oecophoridae)

Plectophila discalis (LEP., Xyloryctidae)

Avaeostoma aenicta (LEP., Xyloryctidae)

Procometis sp. [WARI] (LEP., Xyloryctidae)

Pollamisus viridipulverulentus (LEP., Zygaenidae)
‘Light Brown Apple Moth’ (LEP., ? Epiphyas postvittana,
Tortricidae)

Scoliacma bicolova (LEP., Arctiidae)
Terpna sp. (LEP., Geometridae)
Amisozyga pieroides [QDPI] (LEP., Geometridae)

Doratifera casta, [NSWDA], Doratifera vulnerans [NSWDA],
Doratifera sp. (LEP., Limacodidae)

Anthela aviprepes [NSWDA] (LEP., Anthelidae)

Hyalarcta huebnert, Hyalarcta nigrescens (LEP., Psychidae)

Pseudomantis albofimbriata [NSWDA] (MANT., Mantidae)

Heliothis armigera, Heliothis punctigera (LEP., Noctuidae)

Ipanica cornigera (LEP., Agaristidae)

Acyphyas leucomelas [WARI] (LEP., Lymantriidae)

Pterolocera sp. (LEP., Anthelidae)

Pieris vapae (LEP., Pieridae)

Roeselia metallopa (LEP., Nolidae)

Heliothis punctigera [NSWDA, QDPI], Spodoptera exempta,
Spodoptera mauritia (LEP., Noctuidae)

Unidentified psychid (LEP.)

Anomis planalis [QDPI], Anomis flava [QDPI], Spodoptera
exempta [NSWDA] (LEP., Noctuidae)

Doleschallia bisaltide australis (LEP., Nymphalidae) [Malloch
1929) : 332]

Ialmenus evagovas [QDPI] (LEP., Lycaenidae)

Numerous other LEP. hosts in extra-Australian regions

Animula herrichi [NSWDA], Hyalarcta huebneri (LEP.,
Psychidae)

Leptocneria veducta [NSWDA] (LEP., Lymantriidae)

Loxostege sp. [(QDPI] (LEP., Pyralidae)

Ochrogaster contraria (LEP., Notodontidae)

Phalaenoides glycine [NSWDA] (LEP., Agaristidae)

TACHINIDAE OF AUSTRALIA 175

Spoggosia sp. n.
Stomatomyia tricholygoides

GONIINI

Goniophthalmus australis
Goniophthalmus rufescens

LESKIINI
Apatemyia sp.
Demoticoides pallidus
Sipholeskia sp. ? certima

LEUCOSTOMATINI

Leucostoma simplex

LINNAEMYINI

Chaetophthalmus bicolor
Chaetophthalmus biseriatus

Chaetophthalmus pallipes
Chaetophthalmus spp.

Linnaemya concavicornis
Linnaemya sp.

MINTHOINI
Minthoxia dasyops

NEAERINI

Voriella uniseta

Undescribed gen. & sp. (QLD)
Undetermined genus, sp. n.

NEMORAEINI

Nemoraea sp. n.

PALPOSTOMATINI

Palpostoma aldrichi
Palpostoma desvoidyi
Palpostoma flavum
Palpostoma testaceum

Palpostoma spp.

Anthela excellens (LEP., Anthelidae)

Amata sp. ? aperta [NSWDA] (LEP., Amatidae)

Loxostege affinitalis [NSWDA] (LEP., Pyralidae)

Pseudaletia convecta, Pseudaletia unipuncta [NSWDAI],
Spodoptera exempta (LEP., Noctuidae)

Hehothis armigera, Heliothis sp., Pseudaletia sp. [QDPI],

Spodoptera exempta, Spodoptera mauritia (LEP., Noctuidae)

Neocleptria punctifera [NSWDA], Pandesma quenavadi
(LEP., Noctuidae)

Unidentified curculionid larva (COL.)
‘Cedar Shoot Borer’ (LEP.)
Lygropia clytusalis (LEP., Pyralidae)

Nysius vinitoy (HEM., Lygaeidae)

Agrotis ipsilon, Pseudaletia unipuncta (LEP., Noctuidae)

Hehothis ayrmigerva, Heliothis punctigera, Heliothis sp.
[ODPI], Pseudaletia convecta (LEP., Noctuidae)

Nyctemeva amica, Utetheisa pulchelloides (LEP., Arctiidae)

Euxoa vadians (LEP., Noctuidae)

Agrotis munda, Heliothis punctigera, Heliothis sp.[NSWDA,
QDPI), Persectamia ewingit [NSWDA], Pseudaletia convecta
[NSWDA] (LEP., Noctuidae)

Pseudaletia unipuncta (LEP., Noctuidae)

Maruca testulalis [QDPI} (LEP., Pyralidae)

Barea consignatella (LEP., Oecophoridae)

Cydia molesta [NSWDA},
Tortricidae)

‘Soya Bean Leaf Tier’ (LEP., ? Pyralidae)

Procometis sp. (LEP., Xyloryctidae)

Epiphyas postvittana (LEP.,

Theretva nessus [NSWDA] (LEP., Sphingidae)

Dermolepida albohirta (COL., Scarabaeidae)

Lepidiota frenchi (COL., Scarabaeidae)

Anoplostethus opalinus (COL., Scarabaeidae)

Dermolepida albohivta, Heteronychus arator, Lepidiota
caudata, Metanastes vulgivagus, Pseudholophylla fur-
furacea (COL., Scarabaeidae)

Anomalophylla sp. [Hardy, 1938], Lepidiota trichosterna
[Hardy, 1938], Metanastes vulgivagus, Pseudholophylla
furfuracea (Hardy, 1938]

176

PHASIINI

Alophora aurewentris
Alophora lepidofera
Besserioides varicolor

PROSENINI

Platytainia maculata
Prosena nigripes

RUTILIINI

Amphibolia ignorata

Rutilia s.1. spp.

SIPHONINI

Actia eucosmae
Ceromya norma
Ceromya parviseta

Ceromya sp. ? fergusont

Pervibaea argentifrons

Pervibaea orbata

Peribaea plebeia
Peribaea sp. ? plebeia
Peyibaea sp.

STURMIINI

Anamastax sp. n.
Blepharipa fulviventris

Blepharipa spp.

Eurygastropsis tasmaniae

Palexorista bancrofti
Palexorista macquarti
Palexorista solennis

Palexorista subanajama

K. W.) CROSSKE

Dysdercus sidae (HEM., Pyrrhocoridae)
Nystus vinitoy (HEM., Lygaeidae)
Dysdercus sidae (HEM., Pyrrhocoridae)

Unidentified cerambycid (COL.)
Dermolepida albohirta (COL., Scarabaeidae)

Unidentified melolonthine chafer grubs (COL., Scara-
baeidae) [Paramonov, 1968]

Anoplognathus spp., Dasygnathus sp., Dermolepida sp.,
Lepidiota spp., (COL., Scarabaeidae) [From scattered
literature references: identities of Rutilia species all
suspect]

Crocidosema plebeiana (LEP., Tortricidae)

Pseudaletia convecta [NSWDA] (LEP., Noctuidae)

Isotenes miserana [NSWDA] (LEP., Tortricidae)

Unidentified geometrid (LEP.)

Copromorpha prasinochroa [NSWDA] (LEP., Copromor-
phidae)

Homoeosoma vagella (LEP., Pyralidae)

Acantholeucamia loveyi, Heliothis sp. [QDPI], Pseudaletia
unipuncta, Spodoptera exempta, Spodoptera litura (LEP.
Noctuidae)

Earias huegeli (LEP., Noctuidae)

? Anthela sp. [NSWDA] (LEP., Anthelidae)

Homoeosoma vagella [QDPI] (LEP., Pyralidae)

Panacela lewinae (LEP., Eupterotidae)

Anthela varia (LEP., Anthelidae)

Hippotion celerio, Theretva nessus [NSWDA] (LEP.,
Sphingidae)

Delias argenthona [{QDPI] (LEP., Pieridae)

Papilio aegeus, Papilio anactus, Papilio sp. LEP.,
Papilionidae)

Anthela varia, Anthela sp. (LEP., Anthelidae)

Agyrius convolvuli, Chromis evotus, Hippotion celerio, Theretra
nessus [NSWDA] (LEP., Sphingidae)

Orgyia anartoides (LEP., Lymantriidae)

Plusia sp. (LEP., Noctuidae)

Graphium sarpedon |NSWDA] (LEP., Papilionidae)

Plusia sp. (LEP., Noctuidae)

Unidentified pyralid (LEP.)

Doratifera sp. (LEP., Limacodidae)

Pseudaletia convecta [ODPI], Spodoptera sp. [QDPI] (LEP.,
Noctuidae)

Acantholeucania loveyi (LEP., Noctuidae)

Palexorvista spp.

Paradrino laevicula

Polychaeta nigra
Polychaeta sp.
Sisyvopa sp.
Sturmia convergens

Sturmia sp.
Tritaxys heterocera

Tritaxys milias
Tritaxys sp.
Zygobothria atropivora
Undetermined genus

Undetermined genus
Undetermined genus

TACHININI

Cuphocera emmesia
Cuphocera varia

Microtropesa flaviventris

TRICHOPODINI
Pentatomophaga bicincta

VORIINI
Hyleorus sp.

Hystricovoria sp.
M

TACHINIDAE OF AUSTRALIA 177

A pina callisto (LEP., Agaristidae)

Spilosoma glatignyi (LEP., Arctiidae)

Entometa australasiae (LEP., Lasiocampidae)

Doratifera casta (LEP., Limacodidae)

Habrophylla euryzona (LEP., Lymantriidae)

Heliothis sp. [(QDPI], Persectania ewingui, Plusia sp. [(QDPT],
Pseudaletia convecta, Pseudaletia unipuncta [QDPI],
Spodoptera exempta [NSWDA] (LEP., Noctuidae)

Archernis mitis [NSWDA], Loxostege affinitalis [QDPI),
Maruca testulalis [QDPI] (LEP., Pyralidae)

Unidentified sphingid (LEP.)

Merophyas divulsana [WARI] (LEP., Tortricidae)

Plusia argentifera (LEP., Noctuidae)

Danaus plexippus [QDPI], Euploea core corinna (LEP.,
Nymphalidae)

Delias aganippe (LEP., Pieridae)

Margaronia hyalinata (LEP., Pyralidae)

Unidentified LEP. [NSWDA]

Unidentified LEP. [McFarland Ref. N. 114]

Hymenia vecurvalis [NSWDA] (LEP., Pyralidae)

Brithys crini [QDPI) (LEP., Noctuidae)

Danaus plexippus (LEP., Nymphalidae)

Agyrius convolvuli [ODPI]} (LEP., Sphingidae)

Precis villida (LEP., Nymphalidae)

Acantholeucania loveyi1, Euxoa sp. [NSWDA], Persectania
ewingit, Pseudaletia convecta, Pseudaletia unipuncta (LEP.,
Noctuidae)

Agyrotis infusa [NSWDA], Agrotis ipsilon [QDPI], Euxoa
vadians, Heliothis punctigera [QDPI], Persectania ewingii
[NSWDA] (LEP., Noctuidae)

Pterolocera sp. (LEP., Anthelidae)

Psilogramma menephron (NSWDA] (LEP., Sphingidae)

Numerous other LEP. hosts in extra-Australian regions

Heliocausta rufogrisea [NSWDA] (LEP., Oecophoridae)

Orgyia anartoides [NSWDA] (LEP., Lymantriidae)

Xylorycta luteotactella (LEP., Xyloryctidae)

Unidentified LEP. [McFarland Ref. N. 114]

Acantholeucania loveyi, Pseudaletia unipuncta (LEP.,
Noctuidae)

Also Spodoptera spp. in Oriental Region

Persectania ewingii, Pseudaletia convecta [NSWDA] (LEP.,
Noctuidae)

Amblypelta lutescens (HEM., Coreidae)
Also Axiagastus sp. (HEM., Pentatomidae) in New Guinea

Euproctis lutea (LEP., Lymantriidae)
Unidentified arctiid larva (LEP.) [QDPI]
Aymactica columbina, Xanthodes albago (LEP., Noctuidae)

178 R. W. CROSSKEY

Voria ruralis Plusia argentifera (LEP., Noctuidae)
Numerous other LEP. (also HYM.) hosts in extra-Australian
regions
WINTHEMIINI
Winthemia lateralis Spilosoma glatignyt (LEP., Arctiidae)

Pseudaletia unipuncta [NSWDA] (LEP., Noctuidae)
Winthemia neowinthemioides Brithys crint [(QDPI] (LEP., Noctuidae)
Euploea core corvinna, Euploea sp., Danaus plexippus (LEP.,

Nymphalidae)
Catopsilia pyranthe (LEP., Pieridae)
Winthemia trichopareia Porela arida [NSWDA] (LEP., Lasiocampidae)
Winthemia spp. Anomis flava [QDPI]}, Euxoa sp. [NSWDA], Heliothis

armigera [QDPI], Heliothis sp. [QDPI], Pseudaletia
convecta [NSWDA] (LEP., Noctuidae)
Anaphaeis java [QDPI], Delias aganippe, Delias nigrina
(LEP., Pieridae)
UNDETERMINED TRIBE
Undescribed gen. & sp. Didymuria violescens (PHASM., Phasmatidae)
Tropidoderus childvenii (PHASM., Phasmatidae)
UNDETERMINED TRIBE
Undescribed gen. & sp. Heteronympha merope mevope (LEP., Nymphalidae)
[McFarland Ref. Ny. 33]

HOST-PARASITE LIst

The host orders, and families within each order, are arranged alphabetically.
Host species within each family are arranged in alphabetical order of their valid
binomina, and the author’s name is given for each host species. The tachinid
parasites known for each host are given in alphabetical order of their valid binomina,
the names always corresponding with those considered valid in the taxonomic
catalogue (Part II); subfamily and tribal placements and authors’ names are
omitted for the tachinid parasites as they can all be found easily, if required, from
the taxonomic catalogue, and the tribal positions are clear also from the
‘parasite-host list’ (beginning on p. 171).

The well-known generic instability in the Lepidoptera imposes the need to
provide additional names for the lepidopterous hosts that help to link the
binomina that are currently considered valid with those that actually appear in
literature references or on the data labels attached to reared tachinid specimens.
Whenever necessary earlier generic combinations for the host species are shown
in square brackets on a separate line immediately below the presently valid name.
Similarly, when the specific name that has been in use is now supplanted by a
valid senior synonym the supplanted name is shown in square brackets. In a
very few instances the whole binomen has changed, in which case the whole former
binomen is indicated in square brackets.

Hosts Tachinid Parasites
Order COLEOPTERA
CERAMBYCIDAE
Unidentified larvae Platytainia maculata, Pseudalsomyia pilifacies

TACHINIDAE OF AUSTRALIA

CHRYSOMELIDAE

Aulacophora hilaris Boisduval
Chrysophtharta bimaculata Olivier
Monolepta australis Jacoby

LM. rosea Blackburn]
Paropsis atomaria Olivier

[P. reticulata Marsh]
Paropsis sp.
Unidentified spp.

CURCULIONIDAE

Bryachus squamicollis Pascoe
Gonipterus scutellatus Gyllenhal
Rhabdoscelus obscurus Boisduval

[Rhabdocnemis obscura]
Unidentified larva

SCARABAEIDAE

Anomalophylla sp.
Anoplognathus spp.
Anoplostethus opalinus Brullé
Dasygnathus sp.
Dermolepida albohirta Waterhouse
[Lepidoderma albohivtum]
Heteronychus avatoy Fabricius
[H. sanctaehelenae Blanchard]
Lepidiota caudata Blackburn
Lepidiota frenchi Blackburn
Lepidiota trichosterna Lea
Lepidiota spp.
Metanastes vulgivagus Olliff
(M. blackburm Arrow]
Pseudholophylia furfuracea Burmeister
Unidentified melolonthine larvae

Medinodexia morgant
Paropsivora australis
Monoleptophaga caldwell

Anagonia anguliventris, Froggattimyia
tillyardi, Paropswora grisea

Paropsivora sp.

Anagonia lateralis, Anagonia scutellata

Anagonia lateralis
Anagonia lasiophthalma
Lixophaga sphenophori

Apatemyia sp.

Palpostoma sp.

Rutilia s.l. spp.

Palpostoma flavum

Rutilia s.\. sp.

Palpostoma aldrichi, Palpostoma testaceum
Prosena nigripes

Palpostoma testaceum

?

Palpostoma testaceum

Palpostoma desvoidyi

Palpostoma sp.

Rutilia s.l. spp.

Palpostoma testaceum, Palpostoma sp.

Palpostoma testaceum, Palpostoma sp.
Ampbhibolia ignovata Paramonov

Order DIPTERA

TABANIDAE

Dasybasis hebes Walker
Dasybasis oculata Ricardo

Bactromyiella ? ficta
Bactromyiella ? ficta

Order HEMIPTERA

COREIDAE

Amblypelta lutescens Distant Pentatomophaga bicincta

LYGAEIDAE

Nysius vinitoy Bergroth Alophora lepidofera, Leucostoma simplex

PYRRHOCORIDAE

Dysdercus sidae Montrouzier Alophora aureiventris, Besserioides varicolor

180 R. W. CROSSKEY

Order HYMENOPTERA
ARGIDAE

Zenarge turnert Rohwer

PERGIDAE

Lophyrotoma sp.
[Platypsectra sp. |

Perga dorsalis Leach

Perga glabya Kirby

Pterygophorus analis Costa

Unidentified pergid

VESPIDAE

Polistes tasmaniensis Saussure

Polistes sp.

Polistes sp.

Polistes sp.

Ropalidia marginata jucunda Cameron

Order LEPIDOPTERA
AGARISTIDAE

A pina callisto Walker

Ipanica cornigera Butler

Phalaenoides glycine Lewin
[Aganista g.]

AMATIDAE

Amata ? aperta Walker
[Syntomis ? a.]

ANTHELIDAE

Anthela aviprepes Turner
Anthela excellens Walker
Anthela varia Walker

Anthela sp.
[Darala sp.]
? Anthela sp.

ARCTIIDAE

Nyctemera amica White

Scoliacma bicolova Boisduval
[Scolisoma b.|

Spilosoma glatigny: Le Guilleau

Utetheisa pulchelloides Hampson

Unidentified arctiid

BOMBYCIDAE

Ocinara lewinae Lewin

Zenargomyia moorer

Froggattimyia hirta

Froggattimyia nicholsoni
Froggattimyia hirta
Froggattimyia hirta
Froggattimyia wentwortht

Anacamptomyia nigriventris
Anacamptomyia nigriventyis
Euvespivora decipiens
Koralliomyia sp.
Koralliomyia sp.

Palexorista sp.
Exorista curriei
Exorista sp.

Stomatomyia tricholygoides

Eozenillia vemota

Spoggosia sp. n.

Blephanipa fulviventris, Blepharipa sp., Carcelia
murina

Carcelia murina

Peribaea sp. ? plebeia

Chaetophthalmus biseriatus
Elpe sp., Ethilla translucens

Palexorista sp., Winthemia lateralis

Chaetophthalmus biseriatus
Hyleorus sp.

Elpe sp.

TACHINIDAE OF AUSTRALIA 181

COPROMORPHIDAE
Copromorpha prasinochroa Meyrick

EUPTEROTIDAE

Panacela lewinae Lewin

GELECHIIDAE

Phthorimaea operculeila Zeller
[Gnorimoschema o.|

GEOMETRIDAE

Anisozyga pieroides Walker
[Eucyclodes p.|
Unidentified geometrid

HESPERIIDAE
Hesperilla sp.

LASIOCAMPIDAE

Entometa australasiae Fabricius
[Digglesia a.]|
Porela avida Walker

LIMACODIDAE

Doratifera casta Scott
Doratifera vulnerans Lewin
Doratifera sp.

LYCAENIDAE

Erysichton lineata lineata Murray
Ialmenus evagoras Donovan
Nacaduba biocellata biocellata Felder

LYMANTRIIDAE

Acyphyas leucomelas Walker

Euproctis lutea Fabricius
(Porthesia 1.]

Habrophylla euryzona Lower

Leptocneria veducta Walker
[Lymantria r.]

Orgyia anartoides Walker
[Teta a.]

NOCTUIDAE

Acantholeucama loreyi Duponchel
[Cirphis 1.)

Achaea janata Linnaeus

Agrotis infusa Boisduval

Agyotis ipsilon Hufnagel
[Rhyacia 7.]

Peribaea argentifrons

Anamastax sp. n., Carcelimyia dispar

Trigonospila brevifacies

Ethilla sp.

Ceromya sp. ? ferguson

Chlorotachina sp. n.

Palexorista sp.

Winthemia trichopareia

Austrophorocera sp., Palexorista sp.
Austrophorocera sp., Compsilura concinnata
Austrophorocera sp., Palexorista macquarti

Aplomya sp. ? flavisquama
Carcelia cosmophilae, Exorista sovbillans
Aplomya sp.

Exorista flaviceps
Hyleorus sp.

Palexorista sp.
Exorista sp.

Blephavipa sp., Carcelia sp., undetermined
sturmine genus

Carcelia cosmophilae, Cuphoceravaria, Palexorista
subanajama, Peribaea ovbata, Tritaxys heterocera

Carcelia cosmophilae

Tritaxys milias

Chaetophthalmus bicolor, Tritaxys milias

182

Agrotis munda Walker
Anomis evosa Hiibner
[Cosmophila e.|
Anomis fiava Fabricius
[Cosmophila f.]
Anomis planalis Swinhoe

[A ntarchaea chionosticta Atherton]

Anomis xanthindyma Boisduval
[Cosmophila x.]

Anticarsia tvvovata Fabricius

Armactica columbina Walker

Brithys crint Fabricius

Earias huegeli Rogenhofer
Euxoa vadians Guenée

Euxoa sp.
Heliothis armigera Hiibner
[H. obsoleta misident. ]

Heliothis punctigera Walker

Heliothis spp.

Neocleptria punctifera Walker
Pandesma quenavadi Guenée
Persectania ewingit Westwood

Plusia argentifera Guenée
Plusia sp.

Pseudaletia convecta

Pseudaletia unipuncta Haworth
[Cirphis u.]

Spodoptera exempta Walker
[Laphygma e.|

Spodoptera litura Fabricius
[Prodenia 1.|

Spodoptera mauritia Boisduval

Spodoptera spp.

Xanthodes albago Fabricius
[Acontia malvae Hiibner]

R. W. CROSSKEY

Chaetophthalmus sp.
Carcelia cosmophilae

Carcelia cosmophilae, Carcelia noctuae, Exorista
sovbillans, Winthemia sp.
Exorista sorbillans

Compsilura concinnata

Carcelia noctuae

Hystricovoria sp.

Compsilura concinnata, Sturmia convergens,
Winthemia neowinthemioides

Peribaea plebeia

Carcelia noctuae, Chaetophthalmus pallipes,
Tritaxys milias

Tritaxys hetervocera, Winthemia sp.

Carcelia noctuae, Chaetophthalmus biseriatus,
Exorista curvieit, Goniophthalmus australis,
Winthemia sp.

Carcelia noctuae, Chaetophthalmus biseriatus,
Chaetophthalmus sp., Exorista curriei, Exorista
mungomeryt

Chaetophthalmus biserviatus, Goniophthalmus
australis, Palexorista sp., Peribaea oyrbata,
Winthemia sp.

Goniophthalmus rufescens

Goniophthalmus vufescens

Chaetophthalmus sp., Microtropesa flaviventris,
Palexorista sp., Tritaxys heterocera, Tritaxys
milias

Carcelia cosmophilae, Pavadrino laevicula, Vovria
vyuvalts

Blepharipa sp., Eurygastropsis tasmaniae,
Palexorista sp.

Ceromya norma, Chaetophthalmus biseriatus,
Chaetophthalmus sp., Microtropesa flaviventris,
Palexorista solennis, Palexorista sp., Stomato-
myia tricholygoides, Tritaxys sp., Winthemia
sp.

Carcelia cosmophilae, Carcelia noctuae, Chaeto-
phthalmus bicolor, Cuphocera varia, Linnaemya
concavicornis, Palexorista sp., Pervibaea ovbaia,
Stomatomyia tricholygoides, Tritaxys hetero-
ceva, Winthemia lateralis

Exorista mungomeryi, Exorista sorbillans,
Goniophthalmus australis, Palexorista sp.,
Peribaea orbata, Stomatomyia tricholygoides

Peribaea orbata

Exorista mungomeryi, Goniophthalmus australis
Compsilura concinnata, Palexorista solennis
Hystricovoria sp.

TACHINIDAE OF AUSTRALIA 183

NOLIDAE

Roeselia metallopa Meyrick
[Nola m.]

NOTODONTIDAE

Ochrogaster contraria Walker
Ochrogaster sp.

NYMPHALIDAE

Danaus plexippus Linnaeus
Doleschalha bisaltide australis Felder
Euploea core covinna Macleay

Euploea sp.

Heteronympha merope merope Fabricius
Precis vilida Fabricius

Tisiphone sp.

OECOPHORIDAE

Arachnographa micrastrella Meyrick

Barea consignatella Walker

Heliocausta hemiteles Meyrick
(Garrha h.|

Heliocausta rufogrisea Meyrick
[Garrha r.|

Philobota facialis Fabricius

PAPILIONIDAE

Graphium macleayanus Leech
[Papilio m.]

Graphium sarpedon Linnaeus

Papilio aegeus Donovan

Papilio anactus Macleay

Papilio sp.

PIERIDAE

Anaphaeis java Sparrman
Catopsilia pyvanthe Linnaeus
Delias aganippe Donovan
Delias argenthona Fabricius
Delias nigrina Fabricius
Pieris rapae Linnaeus

PSYCHIDAE

Animula herrichi Westwood
[Thyridopteryx h.]}

Cryptothelea ignobilis Walker
[Clania 7.)

Hyalarcta huebneri Westwood

Hyalarcta nigrescens Doubleday

Metura elongata Saunders
[Otketicus elongatus |

Unidentified psychid

Exorista flaviceps

Carcelimyia dispar, Exorista sp.
Carcelimyia dispar

Pavadrino laevicula, Sturmia convergens

Exorista sorbillans

Payradrino laevicula, Winthemia neowinthemioides
Winthemia neowinthemioides

Undescribed gen. & sp. (? tribe)

Sturmia sp.

Carcelia sp.

Teretvophora fasciata
Minthoxia dasyops
Teretrophora fasciata, Trigonospila brevifacies

Undetermined sturmiine genus

Teretvophora fasciata

Carcelia cosmophilae

Blepharipa sp.

Blepharipa fulviventnis
Blepharipa fulviventris
Blepharipa fulviventris

Winthemia sp.

Winthemia neowinthemioides

Carcelia sp., Pavadrino laevicula, Winthemia sp.
Blepharipa fulviventris

Winthemia sp.

Exorista flaviceps

Exorista sp.

Undetermined exoristine genus
Eozenillia remota, Exorista sp.
Eozenillia remota

Chlorogastropsis orga

Exorista psychidivora

184

PYRALIDAE

Aychernis mitis Turner
Homoeosoma vagella Zeller
Hymenia recurvalis Fabricius
Loxostege affinitalis Lederer
Loxostege sp.

Lygropia clytusalis Walker

[Sylepta c.]

Margaroma hyalinata Linnaeus

[Glyphodes h.]}
Maruca testulahs Geyer
Nacoleia octasema Meyrick
Unidentified pyralids

SPHINGIDAE
Agrius convolvuli Linnaeus
[Herse c.]}
Chromts evotus Cramer
Hippotion celerio Linnaeus

Psilogramma menephron Cramer
[Macrosila casuarinae Walker|

Theretva nessus Drury
Unidentified sphingid
TORTRICIDAE

Aychips australana Lewin
[Cacoecia a.|

Crocidosema plebeiana Zeller

Cydia molesta Busck

Epiphyas postvittana Walker
[Tortrix p.]

Isotenes miserana Walker

Merophyas divulsana Walker

XVEORYCRIDAE

Ayvaeostoma aenicta Turner

Plectophila discalis Walker

Procometis spp.

Xylorycta luteotactella Walker
[Cryptolechia 1.)

ZYGAENIDAE

Pollanisus viridipulverulentus
Guérin-Méneville

UNDETERMINED FAMILY

Order MANTODEA
MANTIDAE

Pseudomantis albofimbriata Stal

Re Wi. CROSSE Y

Palexorista sp.

Peribaea argentifrons, Peribaea sp.
Sisyvopa sp.

Palexorista sp., Stomatomyia tricholygoides
Exorista sp.

Sipholeskia sp. ? cervtima

Pavadrino laevicula

Linnaemya sp., Palexorista sp.
Bactromyiella ficta
Bactromyiella ficta, Palexorista bancrofti

Blepharipa sp., Sturmia convergens

Blepharipa sp.
Blepharipa fulviventris, Blepharipa sp.
Zygobothria atropivora

Blepharipa fulviventris, Blepharipa sp., Nemoraea
sp. n.
Palexorista sp.

Compsilura concinnata

Actia eucosmae
Voriella uniseta
Voriella uniseta

Ceromya parviseta
Palexorista sp.

Undetermined eryciine genus (near Chloro-
gastropsis)

Teretrophora sp.

Undetermined genera (Eryciini and Neaerini)

Undetermined sturmiine genus

Undetermined eryciine genus

Cuphocera emmesia, Polychaeta nigra, Polychaeta

sp.

Exorista covas

TACHINIDAE OF AUSTRALIA 185

Order ORTHOPTERA

ACRIDIDAE
Azelota diversipes Rehn Ceracia fergusont
Chortoicetes terminifera Walker Ceracia fergusont
Coryphistes longipennis Sjostedt Ceracia sp.
Gastrimargus musicus Fabricius Ceracia sp.
Macrotona australis Walker Ceracia fergusont
Urnisa rugosa Saussure Ceracia fergusont

EUMASTACIDAE
Keyacris interpres Rehn Ceracia fergusont
Keyacris marcida Rehn Ceracia fergusont
Moraba amiculi Sj6stedt Ceracia fergusont
Moraba keyi Rehn Ceracia fergusont
Moraba misilliformis Rehn Ceracia fergusont
Moraba viatica Erichson Ceracia fergusoni
Many undescribed spp. Ceracia fergusont

Order PHASMATODEA

PHASMATIDAE
Didymuria violescens Leach Undescribed gen. & sp. (? tribe)
Tropidoderus childrenii Gray Undescribed gen. & sp. (? tribe)

ACKNOWLEDGEMENTS

This work embodies some of the results of a taxonomic study of the Australian
Tachinidae that has proceeded intermittently over the past ten years. During
this period I have been greatly helped by the unstinting generosity of specialist
colleagues in answering my various enquiries or in sending me material, and by
the many non-specialists at other museums and institutions who have so readily
forwarded types to me for study.

It gives me special pleasure to thank my friend Dr Donald Colless (Division of
Entomology, C.S.I.R.O., Canberra) for his constant interest in my work on
Australian Tachinidae and for giving me every possible assistance from the Australian
National Insect Collection.

For the loan of types, other than from Canberra, I warmly thank the following:
Mr C. E. Chadwick (N.S.W. Department of Agriculture, Rydalmere); Dr Willem
Ellis (ZoGlogisch Museum, Amsterdam); Dr P. J. van Helsdingen (Rijksmuseum
van Natuurlijke Historie, Leiden); Professor D. J. Lee (School of Public Health
& Tropical Medicine, Sydney); Dr L. Lyneborg (Universitetets Zoologiske Museum,
Copenhagen); Dr A. Kaltenbach (Naturhistorisches Museum, Vienna); Monsieur
L. Matile (Muséum National d’Histoire Naturelle, Paris); Dr D. K. McAlpine
(Australian Museum, Sydney); Dr G. Morge (Deutsches Entomologisches Institut,
Eberswalde); Dr Per Inge Persson (Naturhistoriska Riksmuseum, Stockholm);
Dr Curtis Sabrosky (U.S.D.A./U.S.N.M., Washington); Dr H. Schumann (Museum
fiir Naturkunde der Humboldt-Universitat, Berlin); Dr D. M. Wood (Entomology
Research Institute, Ottawa); Dr T. E. Woodward (Department of Entomology,

186 R. W. CROSSKEY

University of Queensland, Brisbane); Dr P. Wygodzinsky (American Museum of
Natural History, New York).

I am most grateful to Dr Colless, Professor Lee and Dr McAlpine for the working
facilities and other help given to me at their respective institutions during a visit
to Australia in 1965 and for putting all the tachinid types in their care at my
disposal; and I think Monsieur L. Matile and Dr L. Tsacas for similar assistance
during a visit to the Paris Museum in 1969 to study Macquart’s types.

Dr R. Defretin (Musée d’Histoire Naturelle, Lille) provided me with information
on material in Macquart’s personal collection in Lille, and Dr B. Herting gave me
valuable information on Rondani’s types in Florence, and I gratefully acknowledge
this help. I am indebted to Dr Curtis Sabrosky for providing me with the original
references to the family-group names cited in the catalogue (these are taken from
Dr Sabrosky’s manuscript catalogue of family-group names in the Diptera).

I express my appreciation to the many colleagues, too numerous to name
individually, in the Department of Entomology, British Museum (Natural History)
and the Commonwealth Institute of Entomology who have checked the insect
names cited in the host lists, and have often gone to great pains to divine the
meaning of many of the rather cryptic renderings of host data on the tachinid
data labels.

During the preparation of this work I have taken account of the Tachinidae of
New Guinea collected there personally in 1965. That collecting was supported
financially by the Nuffield Foundation and by the Commonwealth Scientific and
Industrial Research Organization, Canberra, and I am grateful to these bodies for
their assistance.

Finally I thank my wife for writing, on my behalf, voluminous notes on
type-specimens during visits to Australia and Paris.

REFERENCES

[Note : Some works, such as those of Macquart, are better known from their reprint versions
than from the original journals; in these cases the reprint pagination is cited in parentheses
immediately after the journal pagination. }

Acassiz, L. 1846. Nomenclatoris zoologici index universalis, continens nomina systematica
classium, ordinum, familiarvum et generum animalium omnium, tam viventum quam fossilium,
secundum orvdinem alphabeticum unicum disposita, adjectis homonymiis plantarum, nec non
variis adnotationibus emendationbus. Soloduri [=Solothurn, Switzerland], 393 pp.

ALpRICH, J. M. 1922. The Neotropical muscoid genus Mesembrinella Giglio-Tos and other

testaceous muscoid flies. Pvoc. U.S. natn. Mus. 62 (11) : 1-24.

1926. Notes on muscoid flies with retracted hind crossvein, with key and several new

genera and species. Tvans. Am. ent. Soc. 52 : 7-28.

AusTEN, E. E. 1907. The synonymy and generic position of certain species of Muscidae
(sens. lat.) in the collection of the British Museum, described by the late Francis Walker.
Ann. Mag. nat. Hist. (7) 19 : 326-347.

Baranoy, N. 1932. Zur Kenntnis der formosanischen Sturmien (Dipt. Larvaevor.). Neue

Beitr. syst. Insektenk. 5 : 70-82.

1934a. Zur Kenntnis der parasitaren Raupenfliegen der Salomonen, Neubritanniens,
der Admiralitats-Inseln, der Fidschi-Inseln und Neukaledoniens, nebst einer Bestimmung-

stabelle der orientalischen Sturmia-Arten. Vet. Arh. 4 : 472-485.

TACHINIDAE OF AUSTRALIA 187

Baranoy, N., 1934b. Ubersicht der orientalischen Gattungen und Arten des Carcelia-

Komplexes (Diptera : Tachinidae). Tvans. R. ent. Soc. Lond. 82 : 387-408.

1936. Weitere Beitrage zur Kenntnis der parasitaren Raupenfliegen (Tachinidae =

Larvaevoridae) von den Salomonen und Neubritannien. Ann. Mag. nat. Hist.

(10) 17 : 97-113.

1938a. Weiteres iiber die Tachiniden (s.l1.) der Salomon-Inseln. Vet. Arh. 8 : 170-174.

1938b. Neue indo-australische Tachinidae. Bull. ent. Res. 29 : 405-414.

1942. Ein neuer Vespidenparasit von Java und eine mit ihm verwandte Fliege von den

Salomon-Inseln. Vet. Arh. 12 : 161-163.

BECKER, T. 1908. Dipteren der Kanarischen Inseln und der Insel Madeira. Mitt. zool.
Mus. Berl. 4 : 1-206.

BERGROTH, E. 1894. Ueber einige australische Dipteren. Stettin. ent. Ztg 55 : 71-75.

Bezz1, M. 1923. Fissicorn Tachinidae, with description of new forms from Australia and

South America. Proc. Linn. Soc. N.S.W. 48 : 647-659.

1925. On the Tachinid genus Eutherva (Diptera), with description of new species from

Australia, Africa and South America. Proc. Linn. Soc. N.S.W. 50 : 275-283.

1926. A new Tachinid (Dipt.) from Australia, with notes on the forms with obliterated

fourth vein. Ann. Mag. nat. Hist. (9) 17 : 236-241.

1928. Diptera Brachycera and Athevicera of the Fiji Islands based on material in the British

Museum (Natural History). British Museum (Natural History), London, 220 pp.

Bicot, J. M. F. 1874. Diptéres nouveaux ou peu connus. 3° partie, IV. Genres Rutila

et Formosia. Annls Soc. ent. Fr. (5) 4 : 451-467.

1878. Diptéres nouveaux ou peu connus. 9€ partie, XIII. Genres Ocyptera (Latr.),

Ocypterula, Exogaster (Rond.). Annls Soc. ent. Fr. (5) 8 : 40-47.

1880. Diptéres nouveaux ou peu connus. 12° partie, XVIII. Genres Plagiocera

(Macq.), Formosia (Guérin) et Rutilia (Rob.-Desv.). Annls Soc. ent. Fr. (5) 10 : 85-89.

1885a. (Diagnoses de trois genres nouveaux de Diptéres du groupe des Dexiaires).

Annls Soc. ent. Fr. (6) 5 (1885) (Bull. Séanc.) : xi-xii.

1885b. (Diagnoses de deux genres nouveaux de Diptéres du groupe des Dexiaires).

Annls Soc. ent. Fr. (6) 5 (Bull. Séanc.) : xxxii—xxxiii.

1885c. (Diagnoses génériques de deux genres nouveaux de Diptéres du groupe des

Tachinides). Amnnls Soc. ent. Fr. (6) 5 (Bull. Séanc.) : liv—lvi.

1889. Diptéres nouveaux ou peu connus. 34° partie, XLII. Diagnoses de nouvelles

espéces. Annls Soc. ent. Fr. (6) 8 (1888) : 253-270.

Biscuor, J. 1904. Beitrag zur Kenntnis der Muscaria schizometopa. Verh. zool.-bot. Ges.
Wien 54 : 79-101.

Bracxitu, R. E. 1967. A tachinid parasite of Australian grasshoppers. Aust. J. Zool.
15 : 745-758.

Boucut, P. F. 1834. Naturgeschichte der Insekten, besonders in Hinsicht ihrer ersten Zustande
als Larven und Puppen. Lief. 1, 216 pp., Berlin.

Braver, F. 1862. Therobia, eine neue Gattung aus der Familie der Oestriden. Verh.
zool.-bot. Ges. Wien 12 : 1231-1232.

1893. Vorarbeiten zu einer Monographie der Muscaria schizometopa (exclusive Antho-

myidae) von Prof. Dr Fr. Brauer and Julius Edl. v Bergenstamm. Verh. zool.-bot. Ges.

Wien 43 : 447-525.

1899. Beitrage zur Kenntnis der Muscaria schizometopa. Sber. Akad. Wiss. Wien

108 : 495-529.

BRAvER, F. & BERGENSTAMM, J. E. von. 1889. Die Zweifliigler des Kaiserlichen Museums
zu Wien. IV. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive
Anthomyidae). Denkschr. Akad. Wiss., Wien 56 : 69-180 (1-112).

——_ (8 1891. Die Zweifliigler des Kaiserlichen Museums zu Wien. V. Vorarbeiten
zu einer Monographie der Muscaria Schizometopa (exclusive Anthomyidae). Pars II.
Denkschr. Akad. Wiss., Wien 58 : 305-446 (1-142).

188 R. W. CROSSKEY

BRAUER, F. & BERGENSTAMM, J. E. VON. 1893. Die Zweifliigler des Kaiserlichens Museums
zu Wien. VI. Vorarbeiten zu einer Monographie der Muscaria Schizometopa (exclusive
Anthomyidae). Pars III. Denkschr. Akad. Wiss., Wien 60 : 89-240 (1-152).

CotLess, D. H. & McAtrpiIne, D. K. 1970. Diptera (Flies), pp. 656-740 in The Insects of
Austyalia, xiii + 1029 pp., Melbourne University Press.

CoguILLETT, D.W. 1900. Descriptions of two new species of Diptera from Western Australia.
Proc. Linn. Soc. N.S.W. 1900 : 389-390.

1904. New Diptera from India and Australia. Proc. ent. Soc. Wash. 6 : 137-140.

1910. The type-species of the North American genera of Diptera. Pyvoc. U.S. natn.
Mus. 37 : 499-647.

CrossKEY, R. W. 1963. The identity of Tachina convergens Wiedemann, 1824 and Tachina
munda Wiedemann, 1830 (Diptera : Tachinidae). Ann. Mag. nat. Hist. (13) 6 : 77-83.

—— 1964. A new genus and species of Australian Tachinidae (Diptera) parasitic on the

sawfly Zenarge turnert Rohwer (Hymenoptera : Argidae). J. ent. Soc. Qd 3 : 18-22.

1965a. A systematic revision of the Ameniinae (Diptera : Calliphoridae). Bull. Br.

Mus. nat. Hist. (Ent.) 16 : 33-140.

1965b. The immature stages and affinities of the tachinid fly Glaurocara flava, a parasite

of the African bush-cricket Homorocoryphus nitidulus vicinus. Proc. zool. Soc. Lond.

144 : 203-217.

1966a. Generic assignment and synonymy of Wiedemann’s types of Oriental Tachinidae

(Diptera). Ann. Mag. nat. Hist. (13) 8 (1965) : 661-685.

—— 1966b. New generic and specific synonymy in Australian Tachinidae (Diptera). Proc.
R. ent. Soc. Lond. (B) 35 : to1—110 (pagination originally published in error as 95-104).

1966c. The putative fossil genus Palexorista Townsend and its identity with Prosturmia
Townsend (Diptera : Tachinidae). Pyvoc. R. ent. Soc. Lond. (B) 35 : 133-137.

—— 1967a. An index-catalogue of the genus-group names of Oriental and Australasian

Tachinidae (Diptera) and their type-species. Bull. By. Mus. nat. Hist. (Ent.) 20 : 1-39.

1967b. Two new genera and species of eryciine Tachinidae (Diptera) from Australia.

J. Aust. ent. Soc. @ : 27-35.

1967c. A revision of the Oriental species of Palexorista Townsend (Diptera : Tachinidae,

Sturmiini). Bull. Br. Mus. nat. Hist. (Ent.) 21 : 35-97.

1967d. New generic and specific synonymy in Oriental Tachinidae (Diptera). Proc. R.

ent. Soc. Lond. (B) 36 : 95-108.

1969. The type-material of Indonesian Tachinidae (Diptera) in the Zoological Museum,

Amsterdam. Beaufortia 16 : 87-107.

1971. The type-material of Australasian, Oriental and Ethiopian Tachinidae (Diptera)

described by Macquart and Bigot. Bull. Br. Mus. nat. Hist. (Ent.) 25 : 251-305.

1973. A revisionary classification of the Rutiliini (Diptera : Tachinidae), with keys to

the described species. Bull. Br. Mus. nat. Hist. (Ent.) Suppl. 19, 167 pp.

Curran, C. H. 1927a. Three new Tachinidae attacking injurious insects in Queensland.
Bull. ent. Res. 18 : 165-167.

1927b. Some new Australasian and African Diptera of the families Muscidae and

Tachinidae (Dipt.). Ent. Mitt. 16 (5) : 345-357.

1927c. Some new Australasian and African Diptera of the families Muscidae and

Tachinidae (Dipt.). Ent. Mitt. 16 (6) : 438-448.

1929. New Syrphidae and Tachinidae. Ann. ent. Soc. Am. 22 : 489-510.

1930. Four new Diptera from Australia. Am. Mus. Novit. No. 422, 4 pp.

—— 1938a. New species and records of Tachinidae (Diptera). Pyoc. Linn. Soc. N.S.W.

63 : 185-206.

1938b. New Metopiidae and Tachinidae from Africa (Diptera). Am. Mus. Novit.,

No. 985, 8 pp.

Donovan, E. 1805. An epitome of the natural history of the insects of New Holland, New
Zealand, New Guinea, Otaheite, and other islands in the Indian, Southern, and Pacific Oceans :
etc. iv + 41 pls with descriptive text and index (unpaginated), London.

TACHINIDAE OF AUSTRALIA 189

DRaBER-Monko, A. 1965. Monographie der palaarktischen Arten der Gattung Alophora
R.-D. (Diptera, Larvaevoridae). Anmnls zool. Warsz. 23 : 69-194.

DueGpateE, J. S. 1969. A classification of the New Zealand genera of Tachinidae (Diptera :
Cyclorrhapha). N.Z. Jl Sci. 12 : 606-646.

EcceEr, J. 1856. Neue Dipteren-Gattungen und Arten aus der Familie der Tachinarien und
Dexiarien nebst einigen andern dipterologischen Bemerkungen. Verh. zool.-bot. Ver.
Wien 6 : 383-392.

EmpeEn, F. I. van. 1960. Keys to the Ethiopian Tachinidae —III Macquartiinae. Proc.
zool. Soc. Lond. 134 : 313-487.

ENDERLEIN, G. 1936. Klassifikation der Rutiliinen. Verédff. dt. Kolon.-u. Ubersee-Mus.
Bremen 1 : 397-446.

1937. Dipterologica. IV. Sber. Ges. naturf. Freunde Berl. 1937 : 431-443.

Ericuson, W. F. 1842. Beitrag zur Insecten-Fauna von Vandiemensland, mit besonderer
Beriicksichtigung der geographischen Verbreitung der Insecten. Aych. Naturgesch.
1842 (1) : 83-287.

Fasricius, I. C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera,

species, adiectis synonymis, locis, descriptionibus, observationibus. Flensburg & Leipzig,

832 pp.

1794. Entomologia systematica emendata et aucta. Secundum classes, ovdines, genera,
species adjectis synonimis, locis, observationibus, descriptionibus. 4. Copenhagen, 472 +
5 PP:

1805. Systema antliatorum secundum ordines, genera, species adiectis synonymis, locis,

observationibus, descriptionibus. Brunswick, 373 + 30 pp.

FALLEN, C. F. 1810. Fo6rsd6k att bestamma de i Sverige funne Flugarter, som kunna foéras
till Slagtet Tachina. K. svenska VetenskAkad. Handl. [2} 31 : 253-287.

1815. Beskrifning 6fver nagra Rot-fluge Arter, hdrande till slagterna Thereva och
Ocyptera. KX. svenska VetenskAkad. Handl. [3)| 1815 : 229-240.

Gray, G. 1832. In Cuvier, The Animal Kingdom arranged in conformity with its organization
15 (The class Insecta 2), London, 796 pp.

GUERIN-MENEVILLE, F. E. [1831]. Jn Duperrey, ed., Voyage autour du monde sur la corvette

de sa majesté La Coquille, pendant les années 1822, 1823, 1824-1825. Zoologie, Atlas,

Insectes. Paris, 21 plates.

1838. Crustacés, arachnides et insectes. Jn Duperrey, ed., Voyage autour du monde sur
la corvette de sa majesté La Coquille, pendant les années 1822, 1823, 1824-1825. Zool.
Zpt. 2; Div. 1. Paris, 319 pp-

1843. Note monographique sur le genre de Muscides auquel M. Robineau-Desvoidy a
donné le nom de Rutilia, précédée de |’établissement d’un nouveau genre voisin de celui-ci.
Revue zool. 1843 : 262-274.

Harpy, G. H. 1934. Notes on Australian Muscoidea (Calyptrata). Proc. R. Soc. Qd
45 : 30-37.

1938. Notes on Australian Muscoidea III. Dexiinae, Phasiinae, some Tachinidae and
appendix. Proc. R. Soc. Qd 49 : 53-70.

1939. Notes on Australian MuscoideaIV. The genus Microtvopeza and some Phaoniinae.
Proc. R. Soc. Qd 50 : 33-39.

1959. Diptera of Katoomba. Part 3. Stratiomyiidae and Tachinidae. Proc. Linn.
Soc. N.S.W. 84 : 209-217.

HERTING, B. 1962. Neue faunistische und biologische Daten tiber schwedische Tachiniden

(Dipt.). Opusc. ent. 27 : 80-86.

1966. Beitrage zur Kenntnis der europdischen Raupenfliegen (Dipt. Tachinidae).
Stuttg. Beity. Naturk., No. 146, 12 pp.

LE PELeTtier, A. L.M. & SERVILLE, J.G. A. [1828]. In Latreille et al., Encyclopédie méthodique.
Histoive naturelle. Entomologie, ou Histoive naturelle des Crustacés, des Avachnides et des
Insectes 10 (1825) : 345-832.

190 RW. CROSSKEY

Lioy, P. 1864. I Ditteri distribuiti secondo un nuovo metodo di classificazione naturale.
Atti Ist. veneto Sct. (3) 10 : 59-84.

Lorw, H. 1866. Diptera Americae septentrionalis indigena. Berl. ent. Z. 10 : 1-54.

MacguartT, J. 1834. Insectes Diptéres du nord de la France. Athéricéres: Créophiles,

Oestrides, Myopaires, Conopsaires, Scénopiniens, Céphalopsides. Mém. Soc. Sci. Agric.

Lille 1833 : 137-368 (1-232).

1843. Diptéres exotiques nouveaux ou peu connus. 2 (3). Mém. Soc. Sci. Agric.

Lille 1843 : 162-460 (5-304).

1845. Nouvelles observations sur les insectes Diptéres de la tribu des Tachinaires.

Anmnls Soc. ent. Fr. (2) 3 : 237-296.

1846. Diptéres exotiques nouveaux ou peu connus. [1] Supplément. Mém. Soc. Sci.

Agric. Lille 1844 : 133-364 (5-238).

1847. Diptéres exotiques nouveau ou peu connus. 2° Supplément. Mém. Soc. Sci.

Agric. Lille 1846 : 21-120 (5-104).

1848. Diptéres exotiques nouveaux ou peu connus. Suite du 2™¢ Supplément [known

as 3rd Supplement]. Mém. Soc. Sci. Agric. Lille 1847 : 161-237 (1-77).

1851. Diptéres nouveaux ou peu connus. Suite du 4° Supplément publié dans les

Mémoires de 1849. Mém. Soc. Sci. Agric. Lille 1850 : 134-294 (161-364).

1855. Diptéres exotiques nouveaux ou peu connus. 5° Supplement. Mém. Soc. Sci.
Agric. Lille (2) 1 (1854) : 25-156 (5-136).

Mattocu, J. R. 1927. Notes on Australian Diptera. No. xii. Proc. Linn. Soc. N.S.W.

52 : 336-353-

1928a. Notes on Australian Diptera. No. xvii. Proc. Linn. Soc. N.S.W. 53 : 598-617.

— 1928). Notes on Australian Diptera. No. xviii. Proc. Linn. Soc. N.S.W. 53 : 651-662.

— 1929a. Notes on Australian Diptera. No. xix. Proc. Linn. Soc. N.S.W. 54 : 107-117.

—— 1929b. Notes on Australian Diptera. XX. Proc. Linn. Soc. N.S.W. 54 : 283-343.

— 1930a. Notes on Australian Diptera. XXIII. Proc. Linn. Soc. N.S.W. 55 : 92-135.

1930b. Notes on Australian Diptera. XXIV. Proc. Linn. Soc. N.S.W. 55 : 303-353.

— 1930c. Diptera Calyptratae of the Federated Malay States. (Third paper). J. fed.

Malay St. Mus. 16 : 119-153.

1931. Notes on Australian Diptera. XXIX. Proc. Linn. Soc. N.S.W. 56 : 292-298.

1932a. Notes on Australian Diptera. XXXI. Proc. Linn. Soc. N.S.W. 57 : 127-132.

1932b. A new species of Froggattimyia Townsend. Family Tachinidae (Diptera). Aust.

Zool. 7 : 273-274.

1933a. Notes on Australian Diptera. XXXIII. Proc. Linn. Soc. N.S.W. 58 : 74-79.

19330. The Tachinid genus Doddiana Curran (Diptera). Ann. Mag. nat. Hist. (10)

11 : 128-139.

1934. Noteson Australian Diptera. XXXIV. Proc. Linn. Soc. N.S.W.59 : 1-8.

1935. Exotic Muscaridae (Diptera).—XXXIX. (cont.). Ann. Mag. nat. Hist. (10)

16 : 321-343.

1936. Notes on Australian Diptera. XXXV. Pyvoc. Linn. Soc. N.S.W. 61 : 10-26.

1941. Notes on Australian Diptera. XXXIX. Family Chloropidae, Part iii. [Post-

script]. Proc. Linn. Soc. N.S.W. 66 : 41-64.

MEIGEN, J. W. 1800. Nouvelle classification des mouches a deux ailes, (Diptera L.), d’ aprés
un plan tout nouveau. J. J. Fuchs, Paris, 40 pp. [Work suppressed under Opinion 678

of International Commission on Zoological Nomenclature, see Bull. zool. Nom. 20 : 339-342

(1963).]

1803. Versuch einer neuen Gattungs Eintheilung der europaischen zweifliigligen Insekten.

Magazin Insektenk. (Illiger) 2 : 259-281.

1824. Systematische Beschreibung der bekannten Europdischen zweifligeligen Insekten. 4,

xii + 428 pp., Hamm.

MEIJERE, J. C. H. DE. 1917. Studien iiber stidostasiatische Dipteren, XIII. Ueber einige
merkwtirdigen javanischen Dipteren. Tijdschr. Ent. 60 : 238-251.

TACHINIDAE OF AUSTRALIA IQ!

MEsni_, L. P. 10944. Larvaevorinae (Tachininae). Jn Lindner, Fhegen palaearkt. Reg.

64¢ : 1-48.
1947. Révision des Phorocerini de l’ancien monde. (Larvaevoridae). Encycl. ent. (B)
II 10 : 37-80.

1949a. Larvaevorinae (Tachininae). Jm Lindner, Fliegen palaearkt. Reg. 648 : 49-104.

1949). Essai de révision des espéces du genre Drino Robineau-Desvoidy Sturmiinae a

oeufs macrotypes. Bull. Inst. y. Sci. nat. Belg. 25 (42), 38 pp.

—— 1950a. Larvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 648 : 105-160.

1950b. Notes sur les Carceliina (Dipt. Tachinidae) et révision des espéces d’ Afrique.

Revue Zool. Bot. afr. 43 : 1-24.

1951. Larvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 648 : 161-208.

—— 1952. lLarvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 64g : 209-256.

1953. Nouveaux tachinaires d’orient (2° partie}. Bull. Annls Soc. ry. ent. Belg. 89:

146-178.

1954a. Genres Actia Robineau-Desvoidy et voisins (Diptera Brachycera Calyptratae).

Explor. Parc natn. Albert Miss. G. F. de Witte, No. 81, 41 pp.

1954b. Larvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 64¢ : 369-416.

1956. Larvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 648 : 513-560.

— 1960. Larvaevorinae (Tachininae). Jn Lindner, Flegen palaearkt. Reg. 648 : 561-608;

609-656.

1963. Larvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 648 : 801-848.

1965. Note de nomenclature [Dipt. Tachinidae]. Bull. Soc. ent. Fr. 70 : 232.

1966. Larvaevorinae (Tachininae). Jn Lindner, Fliegen palaearkt. Reg. 64g : 881-928.

1968. Nouveaux Tachinaires d’orient (troisiéme série). Bull. Annls Soc. rv. ent. Belg.

104 : 173-188.

Mik, J. 1890. Ugimyia sevicariae Rond., der parasit des japanischen Seidenspinners. Ein

dipterologischer Beitrag. Wien. ent. Zig 9 : 309-3106.

1891. Dipterologische Miscellen. XIX. Wien. ent. Zig 10 : 189-194.

PARAMONOV, S. J. 1950. Notes on Australian Diptera (I-V). Ann. Mag. nat. Hist. (12)
3 2 515-534.

— 1951. Notes on Australian Diptera (VI-VIII). Ann. Mag. nat. Hist. (12) 4 : 745-779.

— 1953. Notes on Australian Diptera (IX—XII). Ann. Mag. nat. Hist. (12) 6 : 195-208.

— 1954. Notes on Australian Diptera (XIII-XV). Ann. Mag. nat. Hist. (12) 7 : 275-297.

1955. Notes on Australian Diptera (XKVI-XIX). Ann. Mag. nat. Hist. (12) 8 : 125-144.

1956. A review of the Australian species of Cylindromyia Meigen and Saralba Walker

(Tachinidae : Diptera). Aust. J]. Zool. 4 : 358-375.

1958. Notes on Australian Diptera (XXVI-XXVIII). Ann. Mag. nat. Hist. (13)

1 : 593-600.

1960. Notes on Australian Diptera (XXIX—XXX). Ann. Mag. nat. Hist. (13) 2

(1959) : 691-704.

1968. A review of the tribe Rutilini (Diptera : Tachinidae). I. Genera other than
Rutilia Robineau-Desvoidy and Foymosia Guérin-Méneville. Aust. J. Zool. 16 : 349-404.

Poxorny, E. 1886. Vier neue 6sterreichische Dipteren. Wuien. ent. Zig 5 : 191-196.

RoBINEAU-DEsvoipy, J. B. 1830. Essai sur les Myodaires. Mém. prés. div. Sav. Acad.
Sct. Inst. Fr. 2 : 1-813.

1848. Myodaires des environs de Paris (suite). Amnnls Soc. ent. Fr. (2) 6 : 429-477.

1851. Myodaires des environs de Paris (suite). Amnls Soc. ent. Fr. (2) 9 : 177-190.

1863. Histoive naturelle des Diptéves des environs de Paris. 1, 1143 pp.; 2, 920 pp.

Paris.

Ronpant, C. 1845. Descrizione di due generi nuovi di insetti Ditteri. Memoria duodecima

per servire alla ditterologia italiana. Nuovi Ann. Sci. nat. Bologna (2) 3 : 25-36.

1856. Genera Italica ordinis Dipterorum ordinatim disposita et distincta et in familias

et stirpes aggregata. Dipterologiae Italicae prodromus 1, 226 pp. [+ 2 p. unnumbered],

Parma.

192 R. W. CROSSKE Y

ROnNDANI, C. 1859. Species Italicae ordinis Dipterorum in genera characteribus definita,
ordinatim collectae, methodo analitica distinctae, et novis vel minus cognitis descriptis.
[Part 2]. Dzipterologiae Italicae prodromus 3, 243 pp. [+ I p. unnumbered], Parma.

1864. Dipterorum species et genera aliqua exotica revisa et annotata novis nonullis
descriptis. Avcho Zool. Anat. Fisiol. 3 (1) (1863) : 1-99. [This work was published in

May 1864 but the reprint version bears the date 1863.]

1865. Diptera Italica non vel minus cognita descripta vel annotata observationibus

nonnullis additis [Fasc. II]. Atti Soc. ital. Sci. nat. 8 : 193-231.

1870. Sull’insetto Ugt. Boll. Soc. ent. ital. 2 : 134-137.

SaBROSKY, C. W. & ARNAUD, P. H. 1965. Family Tachinidae (Larvaevoridae). In Stone
et al. A catalog of the Diptera of America north of Mexico, Agric. Handb. No. 276, U.S.
Department of Agriculture, Washington D.C., 1696 pp.

—— & CrosskKEy, R. W. 1969. The type-material of Tachinidae (Diptera) described by
N. Baranov. Bull. Br. Mus. nat. Hist. (Ent.) 24 : 27-63.

ScCHINER, J. R. 1868. In Reise dey Osterreichischen Fregatte Novara um die Erde inden Jahren
1857, 1858, 1859. Zool. 2 (1), Diptera, 388 pp. Vienna.

Spratt, D. M. & Wo xr, G. 1972. A tachinid parasite of Dasybasis oculata (Ricardo) and
Dasybasis hebes (Walker) (Diptera, Tabanidae). J. Aust. ent. Soc. 11 : 260.

STRAND, E. 1932. Miscellanea nomenclatorica zoologica et palaeontologica. IV. Folia
zool. hydrobiol. 4 : 193-196.

Tuomson, C. G. [1869]. Diptera. Species novas descripsit. In Kongliga svenska fregatten
Eugenies vesa omkving jovden. 2. Zool. 1 (Insecta) : 443-614. K. Svenska Vetenskaps-
Akademien, Stockholm, ‘1868’.

TOWNSEND, C.H.T. 1892a. Notes on North American Tachinidae sens. str. with descriptions

of new genera and species. Paper III. Tvans. Am. ent. Soc. 19 : 88-132.

1892b. The North American genera of Calyptrate Muscidae. Paper III. Tvans. Am.

ent. Soc. 19 : 273-278.

1908. The taxonomy of the Muscoidean flies, including descriptions of new genera and

species. Smithson. misc. Collns 51 (No. 1803) : 1-138.

1912. Areadjustment of muscoid names. Proc. ent. Soc. Wash. 14 : 45-53.
1913. Inquiry into the relationships and taxonomy of the muscoid flies. Can. Ent.

45 : 37-57.

1915a. The family Oestrophasiidae and other notes. Pyvoc. ent. Soc. Wash. 17 : 53-54.
19150. Proposal of new muscoid genera for old species. Proc. biol. Soc. Wash. 28 : 19-24.
1916a. Designations of muscoid genotypes, with new genera and species. Insecutoy

Inscit. mensty. 4 : 4-12.

1916b. Diagnoses of new genera of muscoid flies founded on old species. Proc. U.S.

natn. Mus. 49 (No. 2128) : 617-633.

1916c. New genera and species of Australian Muscoidea. Can. Ent. 48 : 151-160.

1916d. New genera and species of muscoid flies. Proc. U.S. natn. Mus. 51 (No. 2152) :

299-323.

1916e. New muscoid genera (Dip.). Ent. News 27 : 178.

t916f. On Australian Muscoidea, with description of new forms. Imnsecutoy Inscit.

mensty. 4 : 44-45.

1919a. New genera and species of muscoid flies. Proc. U.S. natn. Mus. 56 (No. 2301) :
541-502.
1919b. New muscoid genera, species and synonymy. Imsecutoy Inscit. menstr.

6 (1918) : 157-182.

— 1g1gc. Note on leskiine synonymy (Dipt.). Proc. ent. Soc. Wash. 21: 20.

—— 1921. Some new muscoid genera ancient andrecent. Insecutoy Inscit. mensty.9 : 132-134

1925. Fauna sumatrensis. Calirrhoidae (Dipt. muscoidea). Ent. Mitt. 14 : 250-251.

— 1926a. Fauna sumatrensis. Diptera Muscoidea II. Supplia ent. 14 : 14-42.

—— 1926b. New Holarctic Muscoidea. Insecutor Inscit. menstr. 14 : 24-41.

TACHINIDAE OF AUSTRALIA 193

TownsEND, C. H. T. 1926c. New muscoid flies of the Oriental, Australian, and African
faunas. Philipp. J. Sci. 29 : 529-544.

1927a. Fauna sumatrensis. Diptera Muscoidea III. Swuppita ent. 16 : 56-76.

1927b. Prodiaphania, new name for Diaphania Macquart (1843) preoccupied (Dipt.,

Muscoidea). Ent. News 38 : 159.

1928. New Muscoidea from the Philippines region. Philipp. J. Sci. 34 (1927) : 365-397.

1932. Notes on Old World Oestromuscoid types. Part II. Ann. Mag. nat. Hist.

(10) 9 : 33-57.

1933. New genera and species of Old-World Oestromuscoid flies. Ji N.Y. ent. Soc.

40 (1932) : 439-479.

1934. Five new genera of New Zealand and Malayan Oestroidea. J/ N.Y. ent. Soc.

42 : 247-248.

1936. Manual of Myiology. Part III. 255 pp., Itaquaquecetuba, Sdo Paulo.

1938. Manual of Myiology. Part VII. 434 pp., Itaquaquecetuba, Sao Paulo.

1939. Manual of Myiology. Part VIII. 408 pp., Itaquaquecetuba, Sao Paulo.

— 1940. Manual of Myiology. Part X. 335 pp., Itaquaquecetuba, Sao Paulo.

— 1941. Manual of Myiology. Part XI. 342 pp., Itaquaquecetuba, SAo Paulo.

Tryon, H. 1900. Entomology. Caterpillar plague. Qd agric. J. 6 (2) : 135-147.

VILLENEUVE, J. [1910]. Jn Becker, T., Dipteren aus Siidarabien und von der Insel Sokétra.

Denkschr. Akad. Wiss., Wien 71 (2) : 13i—160.

1911. Description de deux nouveaux Diptéres. Waéen. ent. Ztg 30 : 81-84.

1912. Diptéres nouveaux du Nord Africain [Deuxiéme note]. Bull. Mus. natn. Hist.

nat., Paris 1912 : 505-511.

WALKER, F. 1849. List of the specimens of dipterous insects in the collection of the British

Museum 4 : 689-1172.

1852. Insecta Saundersiana : or characters of undescribed insects in the collection of William

Wilson Saunders, Esq., F.R.S., F.L.S., &c. 1, (Diptera, pt. iv) : 253-414.

1858. Characters of undescribed Diptera in the collection of W. W. Saunders, Esq.,

F.R.S., &c. [Fam. 16. Muscidae]. Tvans. ent. Soc. Lond. (n.s.) [=2nd series], 4 : 190-235.

1859. Catalogue of the dipterous insects collected in the Aru Islands by Mr. A. R. Wallace,

with descriptions of new species. J. Proc. Linn. Soc. Lond. 3 : 77-131.

1861a. Catalogue of the dipterous insects collected at Dorey, New Guinea, by Mr. A. R.

Wallace, with descriptions of new species. J. Proc. Linn. Soc. Lond. 5 : 229-254.

1861b. Catalogue of the dipterous insects collected in Batchian, Kaisaa and Makian,

and at Tidon in Celebes, by Mr. A. R. Wallace, with descriptions of new species. J. Proc.

Linn. Soc. Lond. 5 : 270-303.

1861c. Characters of undescribed Diptera in the collection of W. W. Saunders, Esq.,

F.R.S., &c. Trans. ent. Soc. Lond. (n.s.) [=2nd series] 5 : 268-334. [Pages 268-296 of

this paper were published in 1860 and the remaining pages 297-334 (containing most of

the Tachinidae) in 1861.]

1864. Catalogue of the dipterous insects collected at Waigiou, Mysol, and north Ceram

by Mr. A. R. Wallace, with descriptions of new species. J. Proc. Linn. Soc. Lond.7 : 202-238

1865. Descriptions of new species of the dipterous insects of New Guinea. J. Proc.

Linn. Soc. Lond. 8 : 102-130.

WEstwoop, J. O. 1840. In Westwood, J. O., 1838-1840, Synopsis of the geneva of British
insects. London, 158 pp.

WIEDEMANN, C. R. W. 1824. Munus rectoris in Academia Christiana Albertina aditurus

Analecta entomologica ex Museo Regio Havniensi maxime congesta profert iconibusque

illustrat. Kiel, 60 pp.

1830. Aussereuropdische zweifliigelige Insekten. 2. Hamm, xii + 684 pp.

Wuoutp, F.M.vaANDER. 1881. Diptera medegebracht door de Sumatra-expeditie. Leiden, 60 pp.

1890. In Van der Wulp, 1888-1903. Biologia cent.-am. (Zool.), Insecta, Diptera 2,

489 pp.
1893. Eenige Javaansche Tachininen. Tijdschr. Ent. 36 : 159-188.

N

194 R. We CROSS KEY

ILLUSTRATIONS

vertex ocellar triangle

parafrontal interfrontal area

melee parafrontal

occiput lunula

profrons

face
BA antenna
arista Ae ae
parafacial Sa SS facial
p ridge
epistome
postbucca
vibrissal angle epistome genal dilation
reclinate orbital setae ocellar
seta
inner
vertical
comme 1 seta
proclinate orbital setae
outer
frontal setae vertical

postocular row

vibrissa Peristomal setae
3 RWC

Fics 1-3. Structure and terminology of the head in typical Tachinidae. 1, left lateral
view. 2, facial view. 3, left lateral view showing typical setae. All vestiture omitted
in figs 1 and 2 and hairing omitted in fig. 3.

TACHINIDAE OF AUSTRALIA 195

humeral callus humerals

posthumerals

notopleurals
prescutum

notopleuron

presutural

seta

pre-alar
seta

transverse

Suture 4
supra-alars

scutum F
intra-alars

postalars

postalar callus dorsocentrals

4 acrostichals

basal

lateral

discal

subapical
ie

Fics 4-6. Terminology of thoracic dorsum in Tachinidae. 4, dorsum of thorax, scutellum
omitted, with a complete arrangement of chaetotaxy indicated schematically on one
side only. 5 & 6, two typical shapes of scutellum and terminology of scutellar bristling;
the straight lines connecting basal and subapical setae illustrate important differences
in proportion.

subapical

6 RWC

196 Rk. W. CROSSKE

transverse suture pteropleuron pteropleural seta postalar wall
notopleuron
postalar
scutum callus
suprasquamal
ridge
prescutum scutellum
humeral postscutellum
callus

mediotergite

mesopleuron
pleurotergite

propleuron

propleural seta

prostigmatic seta

posteroventral

declivity of

the thorax
sternopleural setae barette
sternopleuron hypopleuron

N

prosternum

prosternal membrane

edge of fore coxa

8 9 RWG

Fics 7-9. Thoracic characters of Tachinidae. 7, left lateral view of thorax with termin-
ology of sclerites and setae (latter indicated by pore positions only), wing removed.
8, unusual prostigmatic setae in the genus Pevibaea. 9, prothoracic region, drawn from
specimen with setulose prosternum.

TACHINIDAE OF AUSTRALIA 197

mee > ne > F » se Tiole
basal node of Rais second costal sector R343 petiole

Ry

(R445+Myp

m-cu

a E Rand ative i bend of vein M

10

profrons

antennal
axis’ ——

facial
carina

vibrissal
axis
epistomal——

axis

epistome

gena<_

12 B 14

4" 49

RWC

Fics 10-19. Structural characters of Tachinidae. 10, wing of a species with cell R;
open and bend of vein M angulate. 11, apical part of wing of a species with cell R,
closed and bend of vein M rounded. 12, head profile in a typical Prosenine with facial
carina. 13, head profile in a typical Carceliine, showing relative widths of gena and
profrons. 14, head profile showing important landmark axes. 15, showing precoxal
row of hairs on sternopleuron in genus Actia. 16 & 17, ventral view of second costal
sector in (16) forms with the sector bare, and (17) forms with the sector haired. 18 &
19, dorsal view of apex of hind tibia in typical forms (18) with an ad preapical seta,
and (19) without such seta.

198 Re Wie GROSSE

epandrium

St3

St4

surstylus

St5

cercus

21

lobe of St5

UA,

epandrium

basiphallus

epiphallus

surstylus

distiphallus
distiphallus cercus

22 23 RWC

Fics 20-23. Abdomen and ¢ genitalia in representative Tachinidae. 20, ventral view
of abdomen with numbering of tergites (T) and sternites (St). 21, apical view of a
common form of hypopygium, aedeagus and vestiture omitted. 22, ‘L-shaped’ aedeagus
of typical Proseninae. 23, left lateral view of hypopygium in a typical member of the
Goniinae, vestiture omitted.

TACHINIDAE OF AUSTRALIA

Fics 24-31. Head profiles in representative genera of Australian Tachinidae. 24,
Euthera (Q). 25, Gerocyptera (3). 26, Saralba (3). 27, Senostoma (3). 28, Alophora
(subg. Mormonomyia) (9). 29, Ola (3). 30, Platytainia (9). 31, Rutiha (subg.
Chrysorutilia) (3).

199

200 R. W. CROSSKEY

Fics 32-38. Head profiles in representative genera of Australian Tachinidae. 32, Sumpi-

gaster (g). 33, Elpe (3). 34, Doddiana (Q). 35, Thelaiva (3). 36, Halydaia (9). 37,
Voria (2). 38, Nemoraea (9).

TACHINIDAE OF AUSTRALIA 201

Fics 39-46. Head profiles in representative genera of Australian Tachinidae. 39,
Palpostoma (g). 40, Myiotrixa (3). 41, Exechopalpus (3), palp shown. 42, Australo-
tachina (g). 43, Toxocnemis (g). 44, Chaetophthalmus (3). 45, Neximyia (Q). 46,

Zita (g).

202 R. W. CROSSKEY

ee

Nive
Up reese

AWA

Fics 47-53. Head profiles of representative genera of Australian Tachinidae. 47,
Austronilea (g). 48, Carcelimyia (2). 49, Zenargomyia ($). 50, Metaphryno (g). 51,
Palexorista (Q). 52, Koralliomyia (g). 53, Argyrophylax (9).

TACHINIDAE OF AUSTRALIA 203

@----e-----|.,

@ --------------}|-#---«

56

@---e@---e__

61 62 RWC 63

Fics 54-63. Schematic representations of the main patterns of prescutal and scutal
chaetotaxy in Australian Tachinidae. 54, Palpostomatini. 55, Ormiini. 56, Leuco-
stomatini. 57, Glaurocarini. 58, Cylindromyiini. 59, Voriini. 60, Minthoini. 61,
Linnaemyini. 62, Tachinini. 63, Sturmiini. Nofe: arrangement of setae is not
necessarily constant in the tribes indicated, ‘pore’ sizes shown are exaggerated for
clarity, and relative sizes of setae one to another can be gauged approximately from the
sizes of the black circles. Figures based on specimens from following genera: 54, Palpostoma;
55, Lherobia; 56, Leucostoma; 57, Doddiana; 58, Cylindromyia; 59, Hyleorus; 60,
Sumpigaster; 61, Chaetophthalmus ; 62, Cuphocera; 63, Palexorista.

204

R. W. CROSSKEY

70 71 Te RWC

Fics 64-72. The main arrangements of scutellar setae in Australian Tachinidae. 64,

in Alophora (Moymonomyia) (Phasiini). 65, in Palpostomatini. 66, in Alophora
(Hyalomya) (Phasiini). 67, in Siphonini. 68, in Acemyini. 69, in Voriella (Neaerini).
70, in typical Prosenini, drawn from Senostoma. 71, in typical Blondeliini, drawn
from Monoleptophaga. 72, in Carcelia and Winthemia and some other goniine genera
in which subapical setae are unusually widely spaced.

TACHINIDAE OF AUSTRALIA

78

Fics 73-78. Wing venation in representative genera and tribes of Australian Tachinidae.
73, Myiotrixa (Myiotrixini). 74, Pentatomophaga (Trichopodini). 75, Alophora
(Hyalomya) (Phasiini). 76, Cylindromyia (Cylindromyiini). 77, Euthera (Eutherini).
78, Leucostoma (Leucostomatini).

205

206 R. W. CROSSKEY

Fics 79-84. Wing venation in representative genera and tribes of Australian Tachinidae.
79, Prosenina (Prosenini). 80, Pyosena (Prosenini). 81, Doddiana (Glaurocarini). 82,
Hystricovoria (Voriini). 83, Voria (Voriini). 84, Hyleorus (Voriini).

TACHINIDAE OF AUSTRALIA

bo

88 89 RWC

Fics 85-89. Wing venation in representative genera and tribes of Australian Tachinidae.
85, Tvigonospila (Blondeliini). 86, Voriella (Neaerini). 87, Evxorista (Exoristini). 88,
Cevacia (Acemyini). 89, Ceyomya (Siphonini).

~“I

208 RW. CROSSKEY

93 94 95 RWC

Fics 90-95. Abdominal shape in some Australian Tachinidae (vestiture omitted). 90,
Savalba (Trichopodini). 91, Pentatomophaga (Trichopodini). 92, Cylindromyia
(Cylindromyiini). 93, Leucostoma, 9 showing forcipate apex to abdomen. 94, species of
Rutila with excavate T5. 95, Teretrophora, 2 showing exceptionally elongate conical T5.

APPENDIX 209

Two pertinent items of nomenclatural information became known while this
paper was at the page-proof stage. These concern the generic name Spoggosia and
the nominal species Masicera viridiventris.

(1) Spoggosia Rondani. In the present paper this name has been used as valid
for a genus of Exoristini in accordance with the work of Mesnil (1947, 1956, 1960),
but Herting (1972)* has now discovered that the name Chetogena Rondani, 1856,
applies to this genus and should be used as its valid name, since it has priority over
Spoggosia Rondani, 1859. The genus Chetogena Rondani is widespread in Eurasia
and Africa, and has one described and at least one undescribed species in Australia.
The combination Chetogena micropalpis (Malloch, 1930) comb. n. is here established
for the described Australian species. It should be noted that Herting (1972 : 8)
spelt the name as Chaetogena but that the original spelling Chetogena is correct under
the Code.

(2) Masicera viridiventris. Macquart (1847: 84 & 1851: 163) described two
nominal species under this name, the later (1851) use of the name being a junior
primary homonym of the earlier (1847) use. The earlier nominal species was described
from Tasmania and the later one from Egypt. Townsend (1916c) cited the two
viridiventris names as synonyms and stated that the cited provenance ‘Egypte’
for the later use was in error. During earlier work on Macquart’s types (Crosskey,
1971 : 276) I was unable to find the type of viridiventris (2), purportedly from
Egypt, and rejected Townsend’s synonymy of the two viridiventris as unproven,
but it can now be confirmed that Townsend was right. Ona visit to Oxford Univer-
sity Museum in April, 1973, the female holotype of M. viridiventris Macquart, 1851,
was found amongst the Palaearctic Tachinidae from Bigot’s collection, and exam-
ination showed at once that the specimen is undoubtedly conspecific with, and the
female of, M. viridiventris Macquart, 1847 (described from the male); the two types
have been directly compared. The cited provenance of Egypt for vividiventris (2)
is unquestionably in error, and the second use of viridiventris should appear as
follows in the synonymy of Tasmaniomyia viridiventris (Macquart, 1847) on page
152 of this work:

vividiventvis Macquart, 1851: 163 (190) (Masicera). Holotype 9, AUSTRALIA, prob.
TasMaNIA [publ. ‘Egypte’ in error] (UM, Oxford) [examined]. (Name a junior primary
homonym of M. viridiventris Macquart, 1847, no replacement name required.)

The holotype of viridiventris (1851) is in fair condition, but has lost the right mid
leg and the apices of some tarsi, the thoracic dorsum is rather crushed and the
ptilinum partially extruded. It bears Macquart’s original label reading ‘Masicera
viridiventris 9. Macq. n. sp.’ and Bigot’s collection label reading ‘M Viridiventris.
6g. Egypt. Macq.’ (the sex sign being erroneous). At the time of writing the holotype
is still in the collection of the University Museum, Oxford, but it is hoped that
consent will be obtained for it to be transferred to the British Museum (Natural

* HerTING, B. 1972. Die Typenexemplare der von Meigen (1824-1838) beschreibenen Raupenfliegen
(Dipt. Tachinidae). Stuttg. Beitr. Naturk., No. 243, 15 pp.

oO

210 R.W.CROSSKEY

History), where all other types of Tachinidae from Bigot’s extra-Palaearctic material
are housed. (It is germane to note here that on p. 294 of my 1971 paper I mis-
takenly implied that all the Tachinidae from Bigot’s collection are in the BMNH.
This is not so: all the extra-Palaearctic Tachinidae are in the BMNH, but the
Palaearctic material, which includes some Macquart types as well as several Bigot
types of Palaearctic nominal species, is in Oxford.)

INDEX TO FAMILY-GROUP NAMES

The following index is to names of Tachinidae only, host names being excluded.
The main entries for each family-group taxon are indicated by bold type, the first
bold number(s) referring to the treatment in the keys (Part I) and the second

bold number referring to the catalogue entry (Part II). Numbers in italics indicate

the pages on which figures appear.

Acemyidae, 136

Acemyini, 10, 77, 78, 107, 136, 167, 169, 170,
i7lp 172,204,207,

Actiini, 80, 107

Anacamptomyiini, 76, 89, 90-91, 108, 147,
170, I71, 173

Baumhaueriina, 21, 97

Blondelidae, 138

Blondelimi, 10, 11, 13, 15, 22, 23, 24, 25, 53;
75, 77, 81-85, 90, 97, 107, 138, 167, 169,
170, 171, 173, 204, 207

Campylochaetini, 61

Campylochetini, 15, 21, 52, 60-61, 106, 128,
ote 073

Carceliiae, 146

Carceliini, 75, 76, 87, 88, 89-90, 108, 146,
171, 173, 197

Compsilurini, 90

Cylindromyiini, 19, 30, 31, 36-38, 42, 72,
105, 111, 170, 203, 205, 208

Dexiinae, 6, 27, 41, 105, 170
Doleschallini, 13, 17, 42

Echinomyiinae, 50

Eloceriini, 79

Ernestiini, 51, 70-71, 106, 131, 171, 174

Eryciini, 75, 76, 77, 83, 96-99, 108, 154, 156,
169, I7I, 174

Erycinae, 154

Ethillini, 16, 78, 87-88, 89, 107, 144, 169, 170,
eGfiis, 107.A

Ethyllina, 144

Eutherini, 28, 30, 32, 39, 105, 112, 170, 205

Exoristidae, 142

Exoristinae, 75

Exoristini, 11, 78, 85-87, 107, 142, 170, 171,
174, 207

Glaurocarini, 53, 57, 59-60, 106, 127, 171,
203, 206

Gonidae, 136, 153

Goniinae, I0, II, 12, 13, 15, 16, 18, 20, 21, 22,
23, 24, 25, 27, 71, 74-99, 100, 107, 136,
167, 169, 170, 171, 198

Goniini, 24, 60, 75, 76, 96, 108, 153, 171, 175

Gymnosomatini, 30

Helocerini, 79

Leskiini, 10, 44, 51, 67-70, 106, 130, 131,
169, 171, 175

Leucostomatini, 16, 31, 38-39, 105, 112, 170,
175, 203, 205

Leucostomini, I1I2

Linnaemyini, 51, 71, 72-73, 107, 133, 171,
175, 203

Macquartiinae, 50, 106, 171

Masiphyini, 170

Minthoidae, 129

Minithoini, 9, Io, 12, 20, 28, 39, 50, 51, 52,
53, 64-66, 82, 106, 129, 171, 175, 203

Myiotrixini, 51, 55-57, 106, 127, 171, 205

Naereina, 136

Neaerini, 10, 24, 77, 78-80, 107, 136, 171,
175, 204, 207

Nemoraeini, 12, 52, 66-67, 106, 130, 171, 175

Nemoreidae, 130

INDEX 211

Ophirionini, 91

Ormiinae, 127

Ormiini, 13, 15, 17, 25, 27, 50, 57-58, 59,
106, 127, 169, 171, 203

Palpostomatini, 16, 20, 25, 26, 28, 30, 50, 51,
52, 53-55, 106, 126, 169, 171, 175, 203, 204

Palpostomina, 53

Parerigonina, 132

Parerigonini, 51, 71-72, 106, 132, 171

Phasianeae, 109

Phasiidae, 30

Phasimde, 10, II, 13, 15, 19, 20, 21, 22, 23,
25, 27, 28, 29-41, 100, 105, 109, 113, 167,
169, 170

Phasuni, 9, IO, 12, 15, 16, 19, 20, 24, 25, 30,
31, 33-36, 105, 109, 170, 176, 204, 205

Proseninae, 6, 10, II, 13, 15, 17, 20, 22, 23,
25, 27, 28, 41-49, 105, 113, 169, 170, 198

Prosenini, 13, 41, 42, 43-46, 105, 113, 170,
176, 197, 204, 206

Rhinophoridae, 166

Rutiliidae, 118

iting, 6. 10, £2, 13, 16, 17, 19; 21, 22,23,
24, 25, 26, 41, 43, 44, 47-49, 105, 118, 125,
167, 170, 176

Siphonae, 136

Siphonini, 18, 21, 22, 24, 75, 77, 79, 80-81,
107, 136, 171, 176, 204, 207

Strongygastrini, 30

Sturmidae, 148

Sturmiim,) To; "02, 23, 16, 22) 23/124, 75, 76,
82, 87, 88, 91-96, 98, 99, 108, 148, 171,
176, 203

Tachinariae, 109, 126, 134

Tachininaé, ‘50, Fr, 12.03, 15; 20, 20,22, 23;
24, 25, 27, 50-74, 81, 106, 126, 167, 169,
r7r

Tachinini, 24, 51, 72, 73, 74, 107, 134, 171,
177, 203

Thelairini, 23, 44, 52, 63-64, 106, 129, 171

Thelareini, 129

Trichopodini, 31, 32-33, 105, 109, 170, 177,
205, 208

Voriinae, 50
Voriini, 11, 15, 52, 60, 61-63, 106, 128, 171,
177, 203, 206

Winthemiiae, 145
Winthemiini, 16, 75, 76, 87, 88-89, 107, 145,
171, 178

INDEX TO GENUS-GROUP NAMES

The following index is to names of Tachinidae only, host names being excluded.
The main entries for each genus-group taxon are indicated by bold type, the first
bold number referring to the genus (or subgenus) in the keys (Part I) and the second

bold number referring to the catalogue entry (Part IJ).

the pages on which figures appear.

Acemya, 78

Acephana, 138

Actia, 24, 81, 107, 136, 197

Acucera, 44, 46, 105, 113

Acuphocera, 134

Aequia, 112

Afroplagia, 61

Afrovoria, 61, 63

Agalmia, 124

Allophora, 109

Alophora, 6, 12, 20, 24, 28, 30, 33, 34, 35, 36,
105, 109, 110

Alophorella, 33, 35, 109

Alophorophasia, 34

Ameniamima, 48, 121

Numbers in italics indicate

Amphibolia s.l., 5, 19, 25, 48, 105, 118
Amphibolia s.str., 49, 118
Amphibolosia, 133

Amphitropesa, 27, 51, 70, 71, 106, 132
Amplipila, 145

Anacamptomyia, 21, 90, 91, 108, 147
Anagonia, 82, 85, 107, 138, 167
Anamastax, 94, 108, 148, 176
Anaperistommyia, 129

Anatropomyia, 44, 46, 105, 113
Anavoria, 63

Aneogmena, 97, 98

Apalpostoma, 54, 55, 106, 126
Apalpus, 73, 107, 133

Apatemyia, 67, 69, 106, 130, 169, 175

212 INDEX

Apilia, 148

Aplomya, 97, 98, 108, 154, 158, 174
Aprotheca, 99, 108, 133, 154, 158
Archimera, 155

Argyrophylax, 90, 108, 146, 202
Argyrothelaira, 90, 108, 146
Arrhenomyza, 93, 108, 148
Atractodexia, 129

Australotachina, 51, 72, 106, 133, 207
Austrodexia, 43, 117, 158
Austronilea, 97, 99, 108, 155, 202
Austrophasia, 109

Austrophorocera, 85, 87, 107, 142, 174
Austrophryno, 99, 108, 155

Bactromyiella, 76, 83, 97, 108, 155, 171
Ballardia, 133

Besserioides, 34, 35, 105, 110
Bezziomyiobia, 68, 70

Billaea, 13, 44, 46

Blepharella, 22, 82, 94, 108, 148
Blepharipa, 92, 95, 108, 149, 176
Blepharipoda, 149

Bogosia, 32

Bogosiella, 34

Brullaea, 39

Brumptallophora, 34, 110

Calirrhoe, 115

Callotroxis, 41

Calopygidia, 150

Calozenillia, 94, 103, 108, 149

Calyptromyia, 38, 39

Campbellia, 6, 20, 33, 34

Campylochaeta, 61

Campylocheta, 60, 61

Carcelia s.l., 5, 16, 21, 75, 88, 90, 92, 108,
146, 163, 173, 204

Carcelia s.str., 90, 146

Carcelimyia, 89, 90, 108, 147, 202

Ceracia, 21, 78, 107, 136, 172, 207

Ceromya, 81, 107, 137, 158, 207

Ceromyia, 137

Chaetogaster, I19

Chaetogastrina, 118

Chaetolydella, 66

Chaetophthalmus, 5, 12, 18, 22, 71, 73, 107,
113, 175, 201, 203

Chaetoria, 86, 107, 142

Chetogaster, 15, 41, 42, 43, 47, 105, 119

Chlorodexia, 132

Chlorogaster, 152

Chlorogastrina, 152, 158

Chlorogastropsis, 99, 108, 155, 158, 174

Chlorotachina, 70, 71, 106, 132, 174

Chrysopasta, 25, 47, 106, 119

Chrysopygia, 149

Chrysorutilia, 16, 17, 22, 23, 49, 121, 199

Clairvillia, 39

Clara, 36

Clinogaster, 37

Codium, 119

Compsilura, 82, 84, 107, 139

Crossocosmia, 149

Crypsina, 88, 89, 107, 145

Cuphocera, 74, 107, 134, 203

Curtocera, 36

Cylindromyia, II, 15, 18, 20, 25, 31, 36, 37,
38, 105, 111, 203, 205, 208

Cystometopia, 113

Delta, 139

Deltomyza, 84, 107, 139
Demoticoides, 51, 68, 70, 106, 130
Dexia, 41

Dexiomima, 66

Diaphania, 120

Dicephalomyia, 147, 158
Doddiana, 59, 60, 106, 127, 171, 200, 203, 206
Doleschalla, 16, 20, 25
Dolichocolon, 97, 99

Donovanius, 49, 122

Drino, 92, 95

Echinomya, 50

Echrysopasta, 119

Efftayloria, 34, 35, 105, 111
Eipogonoides, 155, 158

Elpe, 17, 60, 61, 106, 128, 173, 200
Eocarcelia, 147, 158
Eocarceliopsis, 147, 158
Eogymnophthalma, 137
Eoparachaeta, 149
Eophyllophila, 82, 83
Eozenillia, 85, 87, 107, 142
Eristaliomyia, 74, 107, 135
Erythrocera, 97, 98, 108, 156
Ethilla, 88, 107, 144, 174
Ethylla, 144

Euamphibolia, 119

Eucarcelia, 147

Euchrysopasta, 119

Eucompsa, 124

Euphasia, 132

Eurygastropsis, 92, 93, 108, 149
Eustacomyia, 55, 106, 126
Eutachina, 142

Euthera, 11, 13, 28, 31,39, 40, 105, 112, r99, 20

INDEX 213

Eutheropsis, 39

Eutrixa, 53

Eutrixoides, 53

Eutrixopsis, 53, 54

Euvespivora, 21, 91, 108, 147

Exechopalpus, 18, 67, 68, 106, 130, 2or

Exorista, 50, 85, 86, 107, 142, 164, 170, 174,
207

Fallenia, 60

Formodexia, 19

Formosia, 13, 16, 19, 41, 47, 106, 119
Frivaldskia, 60, 61

Frivaldszkia, 60

Frivaldzkia, 60

Froggattimyia, 82, 85, 107, 139, 167, 170

Gaediogonia, 154

Geraldia, 44, 46, 105, 113
Gerocyptera, 37, 38, 105, 112, 199
Gerotachina, 135

Glaurocara, 57, 59, 60, 171
Glossosalia, 142
Gonanamastax, 152

Gonia, 17

Goniophana, 92, 152
Goniophania, 152
Goniophthalmus, 96, 108, 153
Grapholostylum, 49, 124
Gymnosoma, 30

Habrota, 121

Halidaya, 64, 129

Halydaia, 21, 52, 63, 64, 106, 129, 200
Hamaxia, 53, 55

Hamaxioides, 54

Hamaxiomima, 53, 55

Hega, 120

Herbstia, 80, 81, 137

Hermya, 36

Heterometopia, 18, 46, 105, 113

Hillia, 85, 143, 160

Hillomyia, 77, 78, 82, 86, 107, 143, 160
Hobartia, 44, 46, 105, 114

Homotrixa, 57

Huttonobesseria, 36

Hyalomya, 33, 36, 110, 204, 205
Hyalomyia, 33, 110

Hyalomyodes, 112, 113

Hyleorus, 52, 61, 62, 106, 128, 177, 203, 206
Hystricovoria, 63, 106, 128, 177, 206

Indosturmia, 149

Koralliomyia, 76, 91, 108, 148, 173, 202
Kosempomyia, 34

Lasiocalypter, 43, 117, 158

Lasiocalyptrina, 43, 117, 158

Lecanipa, 83, 84, 107, 139, 173

Lecanipus, 139

Leiosia, 154, 158

Leucostoma, II, 15, 17, 38, 39, 60, 105, 112,
203, 205, 208

Leverella, 15, 72, 106, 133

Linnaemya, 18, 22, 51, 73, 107, 134, 175

Linnaemyia, 134

Lixophaga, 77, 82, 83, 84, 107, 140

Macreuthera, 39, 112

Macrochloria, 13, 28, 70, 71, 106, 132

Macropia, 129

Macropodexia, 43, 45, 105, 114

Mallochiola, 139

Mauritiodoria, 157

Medinodexia, 83, 107, 140

Megistogastropsis, 65, 66

Menevillea, 123

Mesembriomintho, 130

Metaphryno, 98, 108, 156, 202

Microceromasia, 140

Microrutilia, 49, 124

Microtropesa, 19, 73, 74, 107, 135

Microtropeza, 135

Mintho, 65

Minthodexia, 63

Minthoxia, 52, 65, 106, 129

Monoleptophaga, 83, 107, 140, 204

Mormonomyia, 6, 20, 33, 34, 36, 110, 199, 204

Mycteromyia, 145

Mycteromyiella, 78, 87, 88, 89, 107, 145, 170,
171

Myiotrixa, 13, 27, 53, 55, 106, 127, 201, 205

Myothyria, 78, 136

Neaera, 78

Nemoraea, 16, 66, 106, 130, 175, 200
Nemorilla, 88, 89, 107, 145, 163
Neophasia, 132

Neorutilia, 17, 48, 125

Neuroplagia, 61

Neximyia, 27, 51, 70,.71, 106, 132, 201

Ochromeigenia, 53
Ocyptera, III
Ocypteropsis, III
Oestrocara, 59

Ola, 44, 45, 105, 114, 199

214 INDEX

Opsophana, 138
Opsophasiops, 126
Ormiominda, 58, 127

Pales, 92, 94

Palexorista, 10, 92, 95, 108, 150, 177, 202,
203

Palia, 94, 99, 108, 150

Paliana, 92, 98, 108, 150

Palpostoma, 17, 53, 54, 55, 106, 126, 162,
167, 175, 20L, 203

Parabrachelia, 154, 158

Paradrino, 94, 108, 151

Paragonia, 12, 17, 76, 94, 108, 151

Paramphibolia, 48, 49, 118

Paratropeza, 19, 74

Pareupogona, 83, 84, 107, 140

Paropsivora, 82, 84, 107, 140, 173

Peleteria, 135

Pentatomophaga, 32, 33, 105, 109, 169, 205,
208

Peribaea, 22, 80, 81, 107, 137, 176, 196

Phasioormia, 58

Philotrichostylum, 44, 46

Phorantha, 34, I10

Phorinia, 87

Phorocerosoma, 15, 76, 77, 87, 107, 144, 156,
169, 171

Phorocerostoma, 97, 108, 156

Phryxosturmia, 148

Pilimyia, 84, 107, 141

Plagiprospherysa, 144

Platymya, 155

Platytainia, 44, 46, 105, 114, r99

Podomyia, 148

Podotachina, 142

Pogonagalmia, 124

Polistiopsis, 37

Polychaeta, 92, 93, 108, 151, 177

Preuthera, 39, 40

Prodegeeria, 83

Prodiaphania, 26, 47, 106, 120

Prosena, 13, 43, 44, 45, 105, 115, 206

Prosenina, 44, 45, 105, 116, 206

Prosenostoma, 124

Prosheliomyia, 63

Prosturmia, 150

Protohystricia, 66

Protomeigenia, 139

Protonemoraea, 66

Psaronia, 122

Psaroniella, 125

Psecacera, 44

Pseudalsomyia, 98, 108, 156, 169, 171

Pseudogonia, 96, 108, 154
Pseudokea, 145
Pseudopalpostoma, 126
Pseudotrichopoda, 109
Pygidia, 38, 133
Pygidimyia, 71, 72, 106, 133

Quadra, 96, 108, 151, 153

Redtenbacheria, 40, 41

Rhinomyiobia, 131

Rhinomyobia, 67, 68, 69, 70, 106, 131

Rhynchiodexia, 43, 116, 158

Roederia, 119

Rossimyiops, 167

Rutilia s.l., 16, 25, 47-48, 106, 121, 125, 176,
208

Rutilia s.str., 49, 125

Rutilotrixa, 18, 42, 44, 45, 105, 116, 158

Ruya, 44, 116, 158

Saralba, 32, 33, 105, 109, 199, 208

Schizactiana, 81, 137, 158

Schizactina, 137

Schizoceromyia, 81, 137, 158

Semisuturia, 127

Senometopia, 90, 147, 158

Senostoma, 13, 16, 43, 45, ror, 105, 116, 120,
124, 158, 199, 204

Servillia, 74

Sipholeskia, 67, 68, 69, 106, 131

Siphona, 80

Sisyropa, 92, 94, 96, 108, 151, 177

Spathipalpus, 68

Spoggosia, 21, 75, 86, 97, 107, 143, 175, 209

Steiniomyia, 61, 63

Stenometopia, 147

Stiraulax, 125

Stomatomyia, 18, 75, 85, 86, 107, 144

Stroblomyia, 80, 81, 137

Sturmia, 92, 95, 108, I51, 177

Sumatrodoria, 150

Sumatrosturmia, 149

Sumpigaster, 65, 106, 129, 200, 203

Tachina, 50, 143

Tachinodexia, 65

Talaractia, 137

Tararactia, 137

Tasmaniomyia, 92, 94, 108, 135, 152, 158
Tayloria, III

Teretrophora, 97, 98, 99, 108, 156, 174, 208
Thecocarcelia, 89

Thelaira, 63, 64, 106, 129, 200

INDEX 215

Vespivora, 147
Voria, 52, 61, 62, 106, 128, 164, 200, 206
Voriella, 78, 79, 80, 107, 136, 204, 207

Thelairia, 129
Therairia, 129
Theresia, 44

Therobia, 13, 17, 58, 106, 127, 203
Therobiopsis, 127

Thrycolyga, 142

Thyellina, 92, 153

Tongamyia, 78, 79, 80
Toxocnemis, 67, 69, 106, 131, 20r
Toxonemis, 131

Tricholyga, 143

Trichostylum, 44, 46, 105, 118
Trigonospila, 82, 83, 107, 141, 207
Tritaxys, 5, 92, 93, 108, 152, 177
Tyreomma, 41

Ugimeigenia, 95, 108, 152
Ugimyia, 149
Uschizactia, 137

Vanderwulpella, 38
Verreauxia, 149

Vorina, 142

Winthellia, 92, 94, 108, 153

Winthemia, 10, 23, 88, 89, 92, 107, 145,

204
Winthemya, 145

Xanthodexia, 63
Xanthooestrus, 57
Xenosturmia, 147

Zebromyia, 93, 108, 153
Zenargomyia, 84, 107, 141, 170, 202
Zita, 15, 38, 71, 72, 106, 133, 207
Zoramsceus, 121

Zosteromeigenia, 82, 84, 107, 141
Zosteromyia, 141

Zosteromyiopsis, 141

Zygobothria, 92, 95, 108, 153

INDEX TO SPECIES-GROUP NAMES

178,

The following index is to names of Tachinidae only, host names being excluded.
Names of type-species cited in the catalgoue that are based on extra-Australian

types are omitted except when the species occur in the Australian fauna.

Only

one spelling is given for adjectival specific names whose endings are liable to change
with the generic gender; such names are cited with their currently correct spellings

if considered valid, but with their original spellings if synonyms.

Multiple uses of

the same specific name are distinguished by adding the current genus in brackets.
Numbers printed in bold type indicate entries in the ‘Parasite-host’ list (pp. 172-178)
and therefore show immediately which valid species have known hosts.

accedens, 125

acuminata, 144
acutangulata, 140, 158
aditha, 123, 161

aegyptia, 138

agalmiodes, 123, 124

alata, 34

albicens, 145

albiceps, 145, 158
albifrons, 115, 158, 166
albocincta, 118

albopicta, 124

albovirida, 124

aldrichi, 126, 162, 175, 179
analis (Eurygastropsis), 150

analis (Heterometopia), 114
analoga, 123

anguliventris, 138, 159, 173, 179
angustifrons, 138

angustigena, 122
angustissimifrons, III
anomala, 144

apicale (Palpostoma), 126
apicale (Senostoma), 117, 159
appendiculatum, 117, 158, 159
arcuata, I15

argentata, 115

argentea, 114, 158

argentifera (Chetogaster), 119
argentifera (Rutilia), 121

216 INDEX

argentifera (Sumpigaster), 130
argentifrons, 138, 159, 176, 181, 184
arida, 120

armata, 136, 159

armiceps (Palpostoma), 126
armiceps (Voriella), 79, 157
assimilis, 118

aterrima, III

atratula, III

atripes (Exechopalpus), 131
atripes (Senostoma), 117, 159
atropivora, 95, 153, 177, 184
aurea, 139

aureiventris, 110, 176, 179
aureocauda, 150

aureocincta, 155

aureopyga, 133

auriceps (Chetogaster), 119
auriceps (Exorista), 143

auriceps (Palexorista), 150, 160
auripilis, 149, 159, 166
australasia, 122, 166

australasiae, 113

australiensis, 139

australis (Anamastax), 148
australis (Blepharipa), 149, 158
australis (Doddiana), 128
australis (Goniophthalmus), 153, 175, 182
australis (Paropsivora), 140, 159, 173, 179
australis (Rhinomyobia), 131, 166
australis (Thelaira), 129

australis (Tritaxys), 152

baldwini, 81, 138, 159
bancrofti (Besserioides), 111
bancrofti (Palexorista), 150, 176, 184
bancrofti (Saralba), 109
barcha, 123

basale (Senostoma), 117, 160
basalis (Alophora), I10, 159
basalis (Paliana), 151

bella (Heterometopia), 114
bella (Metaphryno), 156
bella (Prosena), 115
biarmata, 120

bicincta, 32, 109, 177, 179
bicolor (Chaetophthalmus), 134, 166, 175, 181
bicolor (Cylindromyia), 111
bimacula, 111, 158
biserialis, 142

biseriatus, 134, 175, 180, 182
bisetosa (Prosena), 115
bisetosa (Rutilia), 123
braueri (Anamastax), 148

braueri (Therobia), 58

braueri (Trigonospila), 141, 160
brevicornis, 153

brevifacies, 141, 160, 164, 173, 181, 183
brevigaster, 134
brevipalpe, 117, 160, 166
brevis, 137

breviseta, 126
brevisetosa, 145, 158
brevitarsis, 120
brunnea, III

bura, 135, 163

caeruleata, I21

caesia, I2I

caldwelli, 140, 173, 179
calliphon, 97, 157
calliphoroides, 133
calliphorosoma, 132, 158
campbelli (Amphibolia), 118
campbelli (Microtropesa), 135
canberrae (Chetogaster), 119
canberrae (Microtropesa), 135
castanea, 153, 166
castanifrons, 123

castanipes, 123, 125

certima, 67, 131, 160, 175, 184
chersipho, 121

chrysis, 110, 158

ciliata, 153

cilifera, 149

cilipes, 144, 158, 159
cimerascens, 154

cinerea (Carcelia), 147, 163
cinereum (Apalpostoma), 126
cingulata (Rutilia), 121
cingulata (Trigonospila), 141
claripennis, 120
coesiofasciata, 149, 159
commoni (Amphibolia), 118
commoni (Prodiaphania), 120
commune, I17, 160
compressa, 130
concavicornis, 134, 158, 159, 175, 182
concinnata, 139, 173, 181, 182, 184
confluens, 122

confusa (Prosena), 115
confusa (Rutilia), 125

conica, 115, 116, 158
consanguinea, 143, 158
conspicua, 148

convergens, 152, 177, 182, 183, 184
coras, 143, 174, 184

corona, 121

cosmophilae, 147, 173, 181, 182, 183
costalis, 110, 158

crassiseta, 98, 156

cryptica, 122

cupreiventris, 124

curriei, 143, 174, 180, 182
curvipalpis, 150

cygnus, 120

darwini, 137

dasyops, 129, 175, 183
dayi, 111

decipiens, 148, 173, 180
decora, 122

densa, 155

dentata, 125

deserta, 120
despicienda, 157

desvoidyi (Palpostoma), 126, 175, 179

desvoidyi (Rutilia), 125
diadema, 67

dimidiata, 115

discalis, 110, 159

dispar (Carcelimyia), 147, 173, 181, 183

dispar (Prosena), 115
dissimilis, 96, 153

diversa (Ocyptera), 157, 166
diversa (Rutilotrixa), 116, 159
diversa (Winthemia), 145, 158
diversicolor, 155, 156

doddi (Exorista), 143

doddi (Prosena), 115

dorsalis (Apalpus), 133
dorsalis (Prosena), 115, 158
dorsomaculata, 124

dotata, 66

dubitalis, 130

dubitata, 123

durvillei, 125

echinata, 66

echinomides, 120
echinomyidea, 120

elata, 61

elegans (Amphitropesa), 132
elegans (Chrysopasta), 119
elzneri, 152

emmesia, 134, 177, 184
erasmusSi, 114

erecta, 44

erichsoni, 121

erichsonii, 121

erronea, 126

erythropus, 155

INDEX

ethoda, 123
eucosmae, 137, 176, 184
evanescens, 122

faceta, 120

facialis, 156

fallax, 136, 158, 166

fasciata (Teretrophora), 156, 174, 183
fasciata (Trigonospila), 141, 160, 166
fasciatus (Sumpigaster), 130
fenestrata, 111

217

fergusoni (Ceracia), 136, 137, 159, 169, 172,

185

fergusoni (Ceromya), 81, 137, 159, 166, 176,

181
fergusoni (Froggattimyia), 139
fergussoni, 137
ficta, 155, 174, 179, 184
flava (Glaurocara), 59
flaviceps (Chlorotachina), 132, 158

flaviceps (Exorista), 143, 174, 181, 183

flavicornis, 113

flavifrons (Cylindromyia), 111, 158
flavifrons (Doddiana), 128
flavipalpis, 142

flavipes (Apatemyia), 130, 159
flavipes (Rutilia), 124
flavisquama, 154, 159, 174, 181
flavitarsis, 135

flaviventris, 135, 177, 182
flavohirtum, 117, 160

flavum (Palpostoma), 127, 175, 179
formosa, 122

formosina, 122

froggatti, 132

froggattii, 132

frontosa, 121

fulgida, 123,

fullerae, 120

fulvipes (Exechopalpus), 131
fulvipes (Formosia), 120, 166

fulviventris (Blepharipa), 104, 149, 158, 159,

176, 180, 183, 184
fulviventris (Rutilia), 124
funebris, 120
furcata (Prodiaphania), 120
furcatus (Hyleorus), 128
fuscisquama, 124
fuscotestacea, 123

genitalis, 120
georgei, 120
goerlingiana, 122
goniaeformis, 152

218 Ps

graciliseta, 140, 158, 159

grandis, 142

gratiosa, 119

grisea (Anagonia), 138, 159

grisea (Diaphania), 126

grisea (Paropsivora), 140, 173, 179

hardyi, 146

hebes, 155

heterocera, 152, 177, 181, 182
hippobosca, 34, 35, 36, I10, 159
hirta (Eustacomyia), 126

hirta (Froggattimyia), 139, 173, 180
hirticauda, 117, 160

hirticeps (Geraldia), 113
hirticeps (Rutilia), 124
hobartana, III

howeana, III

humeralis, 126

hyalipennis, 157, 166

hyalis, 110, 159

hyria, 117, 160, 165

idesa, 122

ignipennis, 135
ignorata, 118, 176, 179
illota, 147

imitator, 123
imperialis, 122
imperialoides, 122
incomparabilis, 126
inconspicua, 79, 157
inconspicuella, 150
indecisa, 115, 158, 166
inermis, 128

inornata, 123, 125
insectaria, 114
insularis, 58

intensa, I5I
intermedia, 135
inusta, 123

invalida, 137

jacobsoni, 154
jactans, 115
jakovlewii, 25

laevicula, 151, 177, 182, 183, 184
lasiophthalma (Anagonia), 138, 159, 173, 179
lasiophthalma (Pilimyia), 141

lata (Actia), 137

lata (Winthemia), 145, 158

lateralis (Anagonia), 138, 159, 173, 179
lateralis (Blepharella), 149, 158

INDEX

lateralis (Degeeria), 157
lateralis (Pilimyia), 141, 159
lateralis (Rutilotrixa), 116
lateralis (Winthemia), 145, 158, 178, 180, 182
latifrons, III

latigena, 135

latimana, 135

lativentris, 110, 159

lepida, 123

lepidofera, 110, 159, 176, 179
leucosticta, 124

leucozona, 129

limbinevris, 114, 158

lineata, 12I

liris, 124

livida, 155

livis, 124

longicornis, 141, 158, 164
longipes (Apatemyia), 130
longipes (Macropodexia), 114
lucagus, 150

mackerrasi, 129

macquarti, 150, 160, 176, 181
macropus, II5, 158

maculata (Blepharella), 149, 158
maculata (Platytainia), 114, 176, 178
maculiventris, 128

major, 138, 159

malayana, II5

marginalis (Gerocyptera), 37
marginalis (Prosena), 115
marginata, 145, 158

media, 125

melancholica, 146, 159

melas, 157, 165

mellea, 59

micans, 124

micropalpis, 125

milas, 152

milias, 152, 177, 181, 182
mima, 141, 158

minor (Bogosia), 32

minor (Rutilia), 125

minor (Zosteromyia), 141, 166
minuta (Prodiaphania), 120
minuta (Rhinomyiobia), 67, 69
mixtum, I17, 160

modestum, I17, 160
monstruosa, I16, 159
montana, 113

monticola, 138

moorei, 141, 173, 180

morgani, 140, 159, 164, 173, 179

mucrocornis, 157
mungomeryi, 143, 174, 182
murina, 147, 173, 180
mysolana, 144

neowinthemioides, 145, 158, 178, 182, 183
nicholsoni (Froggattimyia), 139, 173, 180
nicholsoni (Prosena), 116

nigra (Polychaeta), 151, 158, 177, 184
nigra (Rutilia), 123

nigribasis, 123

nigriceps, 125

nigricornis, 135

nigricosta, 112

nigrihirta (Alophora), 110, 159
nigrihirtum (Senostoma), 117, 160
nigripalpus, 102, 134, 158, 166
nigripes (Exechopalpus), 131

nigripes (Prosena), 115, 176, 179
nigripes (Rutilia), 125

nigrisquama, 34, 35, 110, 159
nigrithorax, 114

nigriventris, 91, 147, 173, 180
nigrocaerulea, 132

nitens, 122

nitidicauda, 144

nitidiventris, 132, 158, 159

nitidus, 114, 158, 166

noctuae, 147, 173, 182

norma, 137, 176, 182

normalis, T10, 159

norrisi, 129

notatum, 45, 117, 160

obesa, 153,

oblonga (Chetogaster), 119
oblonga (Pareupogona), 140
obtusa, 135

ochriventris, 135
ochromyoides, 111
ocypteroides, 109

olmus, 149

oncoperae, 155

onoba, 123

opaca, 138, 159

orbata, 138, 159, 176, 181, 182
orga, 155, 158, 159, 174, 183
orientalis, 138

ornata (Nemoraea), 66
ornata (Quadra), 151

ornata (Zebromyia), 153

pallens (Doddiana), 128
pallens (Rutilia), 124

INDEX

pallidihirtum, 117, 160
pallidus, 130, 175

pallipes, 134, 175, 182
panthea, 122

paratestacea, 120

parva, 115, 158

parviseta (Ceromya), 137, 176, 184
parviseta (Doddiana), 128
peculiaris, 114

pellucens, 123

pellucida, 119

picta (Calozenillia), 149

picta (Neximyia), 132
pictipennis, 117

pilifacies (Cuphocera), 134, 159
pilifacies (Pseudalsomyia), 156, 174, 178
pilosa, 134, 166

plebeia, 138, 159, 176, 180, 182
plebia, 138

plumifera, 67, 69

polita, 143, 159

portentosa, I51

postulans, 144

potina, 123

prima, 145

profana, 150

prosopina, 55, 127
psychidivora, 143, 174, 183
pubicollis, 122

punctipenne, 117, 160
punctum, 117, 158, 162

quadripunctata, 121
quadriseta, 137
quadrisetosa, 156

recedens, 136

regalis, 123

regina, 120

remota, 87, 142, 159, 174, 180, 183
retusa, 123

rieki (Cylindromyia), 112

rieki (Euthera), 112

rieki (Therobia), 127
rotundatum, 30

rotundipennis, 138
rubricarinatum, 117, 160
rubriceps, 122

rubrifrons, 139

rufescens, 154, 175, 182
tufibarbis, 121

ruficeps, 134, 159

ruficornis (Chlorogastropsis), 155
ruficornis (Rutilia), 122, 125

219

220 INDEX

rufifacies, 139 soror, 126

rufifemur (Cylindromyia), 112 sororcula (Palexorista), 150
rufifemur (Exechopalpus), 131 sororcula (Peribaea), 138
rufifrons, 154 speciosa, 120

rufipalpe (Trichostylum), 45, 118 sphenophori, 82, 140, 160, 167, 173, 179
rufipalpis (Exechopalpus), 131 spinipectus, 123

rufipalpis (Heterometopia), 114, 158 spinolae, 123, 166
rufipalpus (Exechopalpus), 131 splendida, 122

rufipes (Aprotheca), 154 spylosioides, 139

rufipes (Chlorogastropsis), 155, 158, 161 stolida (Amphibolia), 118
rufipes (Macrochloria), 132, 158 stolida (Microtropesa), 135
rufiventris (Alophora), 110, 159 subanajama, 150, 176, 181
rufiventris (Apatemyia), 130 subpubescens, 151, 158
rufiventris (Halydaia), 129 subsessile, 127

rufiventris (Prosena), 115 subtustomentosa, 124
rufofemorata, 131 subvittata, 122
rufolateralis, 133 sumatrensis, 134
rufomaculata, 144, 158 surda, 115

ruralis, 128, 163, 178, 182 sydneyensis, 112

rustica, 164
tasmanensis, 152, 160

sabrata, 123 tasmaniae, 150, 176, 182
salomonica, 148 tasmanica, 89, 146, 165
scutellaris, 146 taylori (Senostoma), 117, 160
scutellata (Anagonia), 85, 139, 159, 173, 179 taylori (Sisyropa), 94, 151
scutellata (Rutilia), 125 tenuis, I16

scutellata (Tritaxys), 152, 160 tenuisetosa, 154, 159

secunda, 127 tessellata (Paropsivora), 140, 159
semifulva, 122 tessellatum (Senostoma), 118, 160
semirufa, 155 testacea (Efftayloria), 111
sensua, 110, 159 testacea (Prodiaphania), 120
serena, 122 testaceicorne, 118, 160

setibasis, 142 testaceipes, 148

setigera (Cuphocera), 134 testaceum (Palpostoma), 126, 127, 162, 166,
setigerum (Senostoma), I17, 160 175, 179

setilatera, 152 tillyardi, 139, 173, 179
setiventre (Phorocerostoma), 156 translucens, 144, 159, 174, 180
setiventre (Senostoma), 117, 160 transversa, 122

setosa, 125 transversalis, 67, 69, 131
sexualis, III, 158 tricholygoides, 144, 175, 180, 182, 184
siberita, 115 trichopareia, 146, 166, 178, 181
sibirita, I15 tricolor, 112

similis (Chaetophthalmus), 134 tristis (Exorista), 143, 159
similis (Masicera), 142 tristis (Gerocyptera), 37, 112
simplex (Leucostoma), 38, 112, 175, 179

simplex (Pareupogona), 140 unguiculata, 112

simplex (Rutilia), 125 unipunctum, 118, 160

sinuata, 135, 136, 166 uniseta, 136, 175, 184

skusei (Euthera), 113, 166 unisetosa, 150

skusei (Microtropesa), 135, 166 uzita, 122

smaragdina, 120

solennis, 150, 176, 182 valentina, 118

solomonica, 146 valida, 81, 137, 159

sorbillans, 143, 174, 181, 182, 183 varia (Cuphocera), 134, 177, 181

s

varia (Prosena), 116, 166
varicolor, 111, 158, 176, 179
variegata (Prosena), 116
variegata (Rutilia), 124
variegatum (Senostoma), 118
varipes (Masicera/Phryno), 156
varipes (Saralba), 109
velutina, 124

versicolor (Chrysopasta), 119
versicolor (Crypsina), 145
vesiculifera, 58

victoriae, 120

vidua, 118, 166

violacea, 119

violacescens, 135, 158, 161
viridescens (Microtropesa), 136
viridescens (Rutilia), 121
viridinigra, 123

viridis, 119

Dr R. W. CrosskeEy, D.Sc., A.R.C.S., F.1I.Biol.

Department of Entomology

British Museum (NaTuRAL History)
CROMWELL Roap

Lonpon SW7 5BD

INDEX 221

viriditestacea, 123
viridithorax, 126
viridiventris, 152, 209
vittata (Cuphocera), 134
vittata (Prodiaphania), 121
vittata (Prosena), 116
vittata (Toxocnemis), 131
vivipara, 125

walkeri, 121

wentworthi, 139, 173, 180
westralica (Cylindromyia), 112
westralica (Rutilotrixa), 116, 159
wilsoni (Amphibolia), 118
wilsoni (Ola), 114

zabirna, 119
zabrina, 119

4M

A LIST OF SUPPLEMENTS
TO THE ENTOMOLOGICAL SERIES
OF THE BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177:

18 plates, 270 text-figures. August, 1965. £4.20.

. SANDS, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera,

Termitidae) from the Ethiopian Region. Pp. 172:500 text-figures. September,
1965. £3.25.

. OxaDA, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso-

philidae. Pp. 129: 328 text-figures. May, 1966. {3.

. GILIOMEE, J. H. Morphology and Taxonomy of Adult Males of the Family

Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967.
£3-15.

. FLETCHER, D. S. A revision of the Ethiopian species and a check list of the

world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146
text-figures,g maps. February, 1967. £3.50.

. HemminG, A. F. The Generic Names of the Butterflies and their type-species

(Lepidoptera: Rhopalocera). Pp. 509. £8.50. Reprinted 1972.

. STEMPFFER, H. The Genera of the African Lycaenidae (Lepidoptera: Rho-

palocera). Pp. 322: 348 text-figures. August, 1967. {8.

. Mounp, L. A. A review of R. S. Bagnall’s Thysanoptera Collections. Pp. 172:

82 text-figures. May, 1968. £4.

. Watson, A. The Taxonomy of the Drepaninae represented in China, with

an account of their world distribution. Pp. 151: 14 plates, 293 text-figures.
November, 1968. {5.

. AFiFI, S. A. Morphology and Taxonomy of Adult Males of the families

Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 51 text-
figures. December, 1968. £5.

. CRosSKEY, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and

its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75.

. Exviot, J.N. An analysis of the Eurasian and Australian Neptini (Lepidoptera:

Nymphalidae). Pp. 155: 3 plates, 1or text-figures. September, 1969. £4.

. GRAHAM, M. W. R. DE V. The Pteromalidae of North-Western Europe

(Hymenoptera: Chalcidoidea). Pp. 908: 686 text-figures. November, 1969.
£19.

. WHALLEY, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198:

68 plates, 15 text-figures. October, 1971. {12.

. SANDS, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae).

Pp. 244: 9 plates, 661 text-figures. July, 1972. {9.90.

. CrossKEY, R. W. A Revisionary Classification of the Rutiliini (Diptera:

Tachinidae), with keys to the described species. Pp. 167: 109 text-figures.
February, 1973. £6.50.

PRINTED BY Unwin Brothers Limited THE GRESHAM PRESS OLD WOKING SURREY ENGLAND

+
ce

ose
ease
=